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Spatial patterns of and competition for resources by territorial carnivores are typically explained by two hypotheses: 1) the territorial defence hypothesis and 2) the searching efficiency hypothesis. According to the territorial defence hypothesis, when food resources are abundant, carnivore densities will be high and home ranges small. In addition, carnivores can maximise their necessary energy intake with minimal territorial defence. At medium resource levels, larger ranges will be needed, and it will become more economically beneficial to defend resources against a lower density of competitors. At low resource levels, carnivore densities will be low and home ranges large, but resources will be too scarce to make it beneficial to defend such large territories. Thus, home range overlap will be minimal at intermediate carnivore densities. According to the searching efficiency hypothesis, there is a cost to knowing a home range. Larger areas are harder to learn and easier to forget, so carnivores constantly need to keep their cognitive map updated by regularly revisiting parts of their home ranges. Consequently, when resources are scarce, carnivores require larger home ranges to acquire sufficient food. These larger home ranges lead to more overlap among individuals' ranges, so that overlap in home ranges is largest when food availability is the lowest. Since conspecific density is low when food availability is low, this hypothesis predicts that overlap is largest when densities are the lowest. We measured home range overlap and used a novel method to compare intraspecific home range overlaps for lionsPanthera leo(n=149) and leopardsPanthera pardus(n=111) in Africa. We estimated home range sizes from telemetry location data and gathered carnivore density data from the literature. Our results did not support the territorial defence hypothesis for either species. Lion prides increased their home range overlap at conspecific lower densities whereas leopards did not. Lion pride changes in overlap were primarily due to increases in group size at lower densities. By contrast, the unique dispersal strategies of leopards led to reduced overlap at lower densities. However, when human-caused mortality was higher, leopards increased their home range overlap. Although lions and leopards are territorial, their territorial behaviour was less important than the acquisition of food in determining their space use. Such information is crucial for the future conservation of these two iconic African carnivores.

Populations of carnivore species outside protected areas may be of considerable importance for conservation, as many protected areas do not provide sufficient space for viable populations. Data on carnivore population sizes and trends are often biased towards protected areas, and few studies have examined the role of unprotected areas for carnivore conservation. We used camera-trapping data and spatial capture–recapture models to estimate population densities for four sympatric carnivores: the African leopard Panthera pardus, spotted hyaena Crocuta crocuta, brown hyaena Parahyaena brunnea and African civet Civettictis civetta in Platjan, a predominantly agricultural, mixed land-use system, South Africa. Mean densities per 100 km2 for the leopard were 2.20 (95% CI 1.32–3.68) and 2.18 (95% CI 1.32–3.61) for left and right flank data, respectively; spotted hyaena, 0.22 (95% CI 0.06–0.81); brown hyaena, 0.74 (95% CI 0.30–1.88); and African civet 3.60 (95% CI 2.34–5.57; left flanks) and 3.71 (95% CI 2.41–5.72; right flanks). Our results indicate that although densities are lower than those reported for protected areas, humans and predators coexist in this unprotected agricultural matrix. We suggest that increased conservation effort should be focused in such areas, to mitigate human–carnivore conflicts. Our study improves the knowledge available for carnivore populations on privately owned, unprotected land, and may benefit conservation planning.

Rodents generate negative consequences for smallholder farmers in Africa that directly impact household and livestock damage, food security and public health. Ecologically Based Rodent Management (EBRM) seeks sustainable solutions for the mitigation of rodent damage through assessments of rodent population dynamics, agro-ecosystems and socio-cultural contexts. We adopt a comparative approach across three rural Afro-Malagasy smallholder farming regions in South Africa, Tanzania and Madagascar to assess the household impacts of rodent pests and current perceptions and preferences associated with several rodent control measures. We conducted focus groups questionnaires and interviews in different study site locations. Rodents assert multiple impacts on Afro-Malagasy farmers demonstrating recurrent and emerging agricultural and household costs, and public health impacts. We identify a significant knowledge gap in educating communities about the application of different EBRM approaches in favour of acute poisons that are perceived to be more effective. Cultural issues and taboos also have a significant impact on the social acceptance of rodent hunting as well as biological control using indigenous predators. We advocate for an enhanced investigation of the socio-cultural beliefs associated with different rodent practices to understand the factors underlying social acceptance. A collaborative approach that integrates the perspectives of target communities to inform the design of EBRM initiatives according to the specific agro-ecosystem and socio-cultural context is necessary to ensure programmatic success.

Theory on intraguild killing (IGK) is central to mammalian carnivore community ecology and top-down ecosystem regulation. Yet, the cryptic nature of IGK hinders empirical evaluations. Using a novel data source - online photographs of interspecific aggression between African carnivores - we revisited existing predictions about the extent and drivers of IGK. Compared with seminal reviews, our constructed IGK network yielded 10 more species and nearly twice as many interactions. The extent of interactions increased 37% when considering intraguild aggression (direct attack) as a precursor of killing events. We show that IGK occurs over a wider range of body-mass ratios than predicted by standing competition-based views, with highly asymmetrical interactions being pervasive. Evidence that large species, particularly hypercarnivore felids, target sympatric carnivores with a wide range of body sizes suggests that current IGK theory is incomplete, underestimating alternative competition pathways and the role of predatory and incidental killing. Our findings reinforce the potential for IGK-mediated cascades in species-rich assemblages and community-wide suppressive effects of large carnivores.

Simple Summary Wildlife ecological patterns are driven not only by environmental and biological contexts, but also by landscape-management schemes that shape those contexts. The present study aims to determine the effect of different environmental factors (including management schemes) on the occurrence patterns of a southern African small mammal community. Based on a landscape where three land-use contexts that differ in their levels of human presence and/or where activities coexist (private ecotourism reserve, mixed farms and traditional communal areas), and by using a body-size-based approach (i.e., using two size-based rodent groups—medium and small—as models), we found that the mean relative abundance of medium-sized species did not differ across the management contexts, but small species’ mean relative abundance was higher in the game reserve. The overall variation in rodent abundance was negatively affected by ungulate presence (possibly linked to a decrease in food availability) and by human presence (increased disturbance). Rodent abundance seems to be influenced by environmental gradients that are directly linked to varying management priorities across land uses, meaning that these communities might not benefit uniformly by the increased amount of habitat promoted by the commercial wildlife industry. Abstract South Africa’s decentralized approach to conservation entails that wildlife outside formally protected areas inhabit complex multi-use landscapes, where private wildlife business (ecotourism and/or hunting) co-exist in a human-dominated landscape matrix. Under decentralized conservation, wildlife is perceived to benefit from increased amount of available habitat, however it is crucial to understand how distinct management priorities and associated landscape modifications impact noncharismatic taxa, such as small mammals. We conducted extensive ink-tracking-tunnel surveys to estimate heterogeneity in rodent distribution and investigate the effect of different environmental factors on abundance patterns of two size-based rodent groups (small- and medium-sized species), across three adjacent management contexts in NE KwaZulu-Natal, South Africa: a private ecotourism game reserve, mixed farms and traditional communal areas (consisting of small clusters of houses interspersed with grazing areas and seminatural vegetation). Our hypotheses were formulated regarding the (1) area typology, (2) vegetation structure, (3) ungulate pressure and (4) human disturbance. Using a boosted-regression-tree approach, we found considerable differences between rodent groups’ abundance and distribution, and the underlying environmental factors. The mean relative abundance of medium-sized species did not differ across the three management contexts, but small species mean relative abundance was higher in the game reserves, confirming an influence of the area typology on their abundance. Variation in rodent relative abundance was negatively correlated with human disturbance and ungulate presence. Rodent abundance seems to be influenced by environmental gradients that are directly linked to varying management priorities across land uses, meaning that these communities might not benefit uniformly by the increased amount of habitat promoted by the commercial wildlife industry.

1.AbstractTraditional cultural beliefs influence perceptions of animals, and in some cases can result in persecution of wildlife. Stigmas against species associated with witchcraft, for example, can act as a barrier to the uptake of more cost effective, sustainable, and environmentally sound practices such as reducing crop damage by controlling rodent agricultural pests by relying on indigenous predators rather than pesticides. One way of enhancing perceptions of wildlife to increase participation in such ecologically-based rodent management (EBRM) schemes, is the development of environmental education initiatives. Low intensity programmes are cost-effective and can produce positive attitudinal shifts, but their impact has not been assessed for species strongly associated with witchcraft. We set out to test whether a single presentation on the natural history of owls (order Strigiformes) could improve perceptions of these species, and increase willingness to participate in an EBRM scheme that involved the installation of owl boxes to increase owl populations and reduce rodent populations and crop damage in agricultural fields. We used a questionnaire survey to assess perceptions of owls at four schools in two villages in South Africa. Our initial survey sampled perceptions of respondents before listening to the presentation. A follow-up survey conducted three months later sampled the perceptions of respondents that had listened to the presentation as well as perceptions of a control group that did not listen to the presentation. We found that associations between owls and witchcraft was a common theme driving negative perceptions of owls. Respondents that watched the presentation had more positive perceptions of owls than respondents that had not watched the presentation, and they were more likely to be willing to put up owl boxes near their home. Despite this shift, negative perceptions of owls still dominated responses due to cultural associations with the occult. These findings indicate that even low-intensity programmes can be effective at enhancing perceptions of wildlife associated with witchcraft. We suggest that environmental education programmes featuring culturally taboo species should adopt a culturally sensitive and locally tailored approach, focus on the benefits these species provide, and may be more effective when delivered with greater intensity.

This work was supported by the Peace Parks Foundation; G.C.S. was funded by a doctoral grant from Fundacão para a Ciência e a Tecnologia (FCT: PD/BD/114037/2015); L.H.S. was supported by the National Research Foundation, South Africa (UID: 107099 and 115040) and by the African Institute for Conservation Ecology. Apex predator reintroductions have proliferated across southern Africa, yet their ecological effects and proposed umbrella benefits of associated management lack empirical evaluations. Despite a rich theory on top-down ecosystem regulation via mesopredator suppression, a knowledge gap exists relating to the influence of lions (Panthera leo) over Africa's diverse mesocarnivore (less than 20 kg) communities. We investigate how geographical variation in mesocarnivore community richness and occupancy across South African reserves is associated with the presence of lions. An interesting duality emerged: lion reserves held more mesocarnivore-rich communities, yet mesocarnivore occupancy rates and evenness-weighted diversity were lower in the presence of lions. Human population density in the reserve surroundings had a similarly ubiquitous negative effect on mesocarnivore occupancy. The positive association between species richness and lion presence corroborated the umbrella species concept but translated into small differences in community size. Distributional contractions of mesocarnivore species within lion reserves, and potentially corresponding numerical reductions, suggest within-community mesopredator suppression by lions, probably as a result of lethal encounters and responses to a landscape of fear. Our findings offer empirical support for the theoretical understanding of processes underpinning carnivore community assembly and are of conservation relevance under current large-predator orientated management and conservation paradigms. Postprint

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Genetic diversity within partial 18S rRNA sequences from Hepatozoon protozoan parasites from wild felines in South Africa was assessed and compared with data from domestic cats to assess patterns of host specificity. Lions, leopards, servals, a caracal and an African wildcat were all positive for parasites of the Hepatozoon felis-complex. However, haplotypes were not species-specific, and potential mixed infections were widespread. Additional genetic markers are needed to untangle the extremely complex situation of these parasites in both domestic cats and wild felines in South Africa.

One of the key factors influencing the population dynamics of threatened species such as felids is disease, but long-term studies of the factors influencing seroprevalence of wild felids are extremely rare, hindering conservation efforts. We set out to determine seroprevalence of six viral diseases (feline panleukopenia virus, feline leukemia virus, feline coronavirus, feline calicivirus, feline herpes virus, and feline immunodeficiency virus) among a population of serval (Leptailurus serval) with an extremely high density in South Africa. We captured 55 individuals over four years and screened blood samples for antibodies to each virus. We found that seroprevalence were high (ranging from 30.0% positive for a single virus to 1.8% positive for up to five viruses) and that seroprevalence was influenced by season and sex, but not body condition. We suggest further monitoring of this population and recommend that long-term studies are conducted for serval and other felids to determine whether these trends are representative on a broader scale. Supplementary Information The online version contains supplementary material available at 10.1007/s10393-021-01533-z.

Conservation agriculture (CA), with reduced tillage, permanent soil cover and diversified cropping systems, is advocated in southern Africa to improve soil quality, reduce input costs and mitigate climate-induced risks. However, improvements in terms of yield and soil organic carbon (SOC) under CA are slow and variable and many small-scale farmers are unable to buffer themselves against potential short-term financial losses. In this study we examined the effects of CA-related management practices on SOC sequestration and productivity at two medium-term sites on a sandy soil (eight year trial) and clay soil (six years) in maize producing areas of South Africa. Using field data, current input costs and market prices for crops, we calculated the gross margin for each system. Treatments compared conventional ploughing under maize monoculture with reduced tillage, intercropping and crop rotation. On the clay soil, SOC was increased under reduced tillage (57.6 t C ha−1) compared to conventional tillage (54.9 t C ha−1) while there was no difference for the sandy soil (19.7 t C ha−1 average across treatments). Profitability was most strongly influenced by seasonal rainfall, but was higher on the sandy soil than the clay soil, with an average gross margin of R11,344 ha−1 and R5,686 ha−1, respectively. This study has demonstrated that while certain CA practices can create site-specific benefits for farmers, it is highly dependent on local weather and soil conditions. For the clay soil an additional payment scheme would be required to reward farmers in southern Africa for C-sequestration to make CA profitable and achieve increased C-mitigation through soil sequestration.

Africa is endowed with a diverse guild of small carnivores, which could benefit stakeholders by providing ecosystem services while fostering conservation tolerance for carnivores. To investigate the potential of small carnivores for the biological control of rodents within agro-ecosystems, we assessed both the ecological and social landscapes within two rural villages in the Vhembe Biosphere Reserve, South Africa. We employed a camera trapping survey underpinned by an occupancy modelling framework to distinguish between ecological and observation processes affecting small carnivore occupancy. We also used questionnaires to investigate perceptions of small carnivores and their role in pest control. We found the greatest diversity of small carnivores in land used for cropping in comparison to grazing or settlements. Probability of use by small carnivores was influenced negatively by the relative abundance of domestic dogs and positively by the relative abundance of livestock. Greater carnivore diversity and probability of use could be mediated through habitat heterogeneity, food abundance, or reduced competition from domestic carnivores. Village residents failed to appreciate the role of small carnivores in rodent control. Our results suggest that there is significant, although undervalued, potential for small carnivores to provide ecosystem services in agro-ecosystems.

Conservation agriculture (CA) is advocated as a sustainable farming method to improve soil health, increase crop yields and food security, while reducing input costs. In South Africa, a country wit...

Intra-guild interactions related to facilitation and competition can be strong forces structuring ecological communities and have been suggested as particularly prominent for large carnivores. The African lion (Panthera leo) is generally thought to be a dominant predator where it occurs and can be expected to have broad effects on sympatric carnivore communities. We used data from two small game reserves in northern South Africa to relate the presence of African lions to abundance, habitat use, diet, and prey selection of two sympatric large carnivores, brown hyaenas (Parahyaena brunnea) and leopards (Panthera pardus). Our results offered some support for the facilitative effects of lions on brown hyaenas, and competitive effects on leopards. However, differences between populations living without and with lions were restricted to broad diet composition and appear not to have permeated into differences in either prey selection, abundance or habitat use. Therefore, we suggest that the potential effects of lions on the predator–prey interactions of sympatric predators may have been context dependent or absent, and subsequently argue that lions may not necessarily influence the predator–prey dynamics in the landscapes they live in beyond those caused by their own predatory behaviour., This research was funded by the National Geographic/Wait’s Foundation (grant number W32-08 to FD), the National Research Foundation in South Africa (grant numbers SFP2008072900003 to FD, IFR2011032400087 and UID 115040 to LS), and the Ministry of Economy and Competitiveness in Spain (grant number RYC2013-14662 to FD).

Understanding resource selection and quantifying habitat connectivity are fundamental to conservation planning for both land-use and species management plans. However, datasets available to management authorities for resource selection and connectivity analyses are often highly limited and fragmentary. As a result, measuring connectivity is challenging, and often poorly integrated within conservation planning and wildlife management. To exacerbate the challenge, scale-dependent resource use makes inference across scales problematic, resource use is often modelled in areas where the species is not present, and connectivity is typically measured using a source-to-sink approach, erroneously assuming animals possess predefined destinations. Here, we used a large carnivore, the leopard Panthera pardus, to characterise resource use and landscape connectivity across a vast, biodiverse region of southern Africa. Using a range of datasets to counter data deficiencies inherent in carnivore management, we overcame methodological limitations by employing occupancy modelling and resource selection functions across three orders of selection, and estimated landscape-scale habitat connectivity – independent of a priori source and sink locations – using circuit theory. We evaluated whether occupancy modelling on its own was capable of accurately informing habitat connectivity, and identified conservation priorities necessary for applied management. We detected markedly different scale-dependent relationships across all selection orders. Our multi-data, multi-scale approach accurately predicted resource use across multiple scales and demonstrates how management authorities can more suitably utilise fragmentary datasets. We further developed an unbiased landscape-scale depiction of habitat connectivity, and identified key linkages in need of targeted management. We did not find support for the use of occupancy modelling as a proxy for landscape-scale habitat connectivity and further caution its use within a management context. Synthesis and applications. Maintaining habitat connectivity remains a fundamental component of wildlife management and conservation, yet data to inform these biological and ecological processes are often scarce. We present a robust approach that incorporates multi-scale fragmentary datasets (e.g. mortality data, permit data, sightings data), routinely collected by management authorities, to inform wildlife management and land-use planning. We recommend that management authorities employ a multi-data, multi-scale connectivity approach—as we present here—to identify management units at risk of low connectivity.

Frugivory and seed dispersal contribute to the maintenance and regeneration of plant communities through transportation of seeds and enhancing germination through seed processing mechanisms. The effects of mammalian frugivore seed processing mechanisms on seed germination are generally well studied and the potential benefits include disinhibition (pulp removal), scarification (gut passage) and fertilisation (from faecal matrix). Nevertheless, our review found that there is bias in the comparative treatments included in seed dispersal studies through exclusion of entire fruit control groups and the fertiliser effect. In this study, we aimed to address such bias by using ecologically relevant experiments to investigate the influence of seed processing mechanisms on germination probability and latency of selected locally abundant fleshy-fruiting plant species, common in the diet of samango monkeys (Cercopithecus albogularis schwarzi), and who are seed-spitters (disinhibition) and seed-swallowers (gut-passage and fertiliser effects). We designed experiments to isolate the cumulative effects of seed processing mechanisms and tested the effects of five treatments and one control treatment (entire fruit). We further assessed if exclusion of ecologically relevant seed treatments or relevant controls would affect our interpretation of the impact of the disperser on seed germination. Comparing gut passage and disinhibition indicated negative effects, whereas comparing gut passage and entire fruit controls indicated neutral effects in one species. Compared with gut passage alone, the fertiliser effects indicated positive or neutral benefits on germination probability. Our study demonstrated that the impacts of mammalian frugivores on germination may be under- or over-estimated in ecological literature where relevant treatments and meaningful controls are excluded.

Due to the strong individual cost of being predated, potential prey species alter their behavior and physiology in response to predation risk. Such alterations may cause major indirect consequences on prey populations that are additive to the direct demographic effects caused by prey being killed. However, although earlier studies showed strong general effects of the presence of apex predators, recent data suggest that indirect effects may be highly context dependent and not consistently present. We combined behavioral data with data on endocrine stress and stable isotopes to assess landscape level effects of lion (Panthera leo) presence on two prey species in South Africa, impala (Aepyceros melampus) and blue wildebeest (Connochaetes taurinus). We also evaluated if there was any seasonal variation in such effects. In addition, we provide results from a physiological validation for an enzyme-linked immunoassay (EIA) that can be used for non-invasive monitoring of glucocorticoid stress metabolite concentrations in impala from fecal pellets. We did not find any significant differences in vigilance behavior, fecal glucocorticoid metabolite concentrations, δ13C values or isotope niche breadth between animals living with and without lions for either species. However, wildebeest living in a reserve with lions spent more time foraging compared to wildebeest in a lion-free environment, but only during the wet season. Values of fecal δ15N suggest a shift in habitat use, with impala and wildebeest living with lions potentially feeding in less productive areas compared to animals living without lions. For both species, characteristics of the social groups appeared to be more important than individual characteristics for both foraging and vigilance behavior. Our results highlight that antipredator responses may be highly dynamic and scale-dependent. We urge for further studies that quantify at what temporal and spatial scales predation risk is causing indirect effects on prey populations.

As the environment becomes increasingly altered by human development, the importance of understanding the ways in which wildlife interact with modified landscapes is becoming clear. Areas such as industrial sites are sometimes presumed to have little conservation value, but many of these sites have areas of less disturbed habitats around their core infrastructure, which could provide ideal conditions to support some species, such as mesocarnivores. We conducted the first assessments of the density of serval (Leptailurus serval) at the Secunda Synfuels Operations plant, South Africa. We ran three camera trap surveys to estimate serval density using a spatially explicit capture recapture framework. Servals occurred at densities of 76.20-101.21 animals per 100 km2, which are the highest recorded densities for this species, presumably due to high abundance of prey and the absence of persecution and/or competitor species. Our findings highlight the significant conservation potential of industrialised sites, and we suggest that such sites could help contribute towards meeting conservation goals.

Tissue samples from wildlife from South Africa were opportunistically collected and screened for haemoprotozoan parasites using nonspecific PCR primers. Samples of 127 individuals were tested, comprising over 50 different species. Haemogregarines were the most commonly identified parasites, but sarcocystids and piroplasmids were also detected. Phylogenetic analyses estimated from the 18S rDNA marker highlighted the occurrence of several novel parasite forms and the detection of parasites in novel hosts. Phylogenetic relationships, which have been recently reviewed, appear to be much more complex than previously considered. Our study highlights the high diversity of parasites circulating in wildlife in this biodiverse region, and the need for further studies to resolve taxonomic issues.

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 the number of owls killed on the roads, 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.

Conservation agriculture (CA) is advocated as a management system for sustainable productivity, while preserving the environment simultaneously. CA has many advantages, but weed management is regarded as one of its biggest challenges. This study reports on the temporal variation in weed occurrence and biomass under conservation and conventional farming practices. The treatment design was a split-split plot, with a randomised complete block design with three blocks as replicates. Tillage was the main plot factor (reduced tillage [RT] and conventional tillage [CT]), and treatments (a combination of cropping systems and fertilizer levels) were treated as the sub-plot factor. Only cultivation year (F(2.48) = 9.12, p < 0.001) and the cultivation year and tillage interaction (F(2.48) = 22.41, p < 0.001) significantly affected weed biomass. Weed biomass and species diversity increased under RT from cultivation year 3 to 5. Under CT weed biomass had a slight downward trend and species composition was similar across the three years with two dominant weeds representing between 87.2 and 75.1% of total weed biomass. The results suggest that tillage practices can affect both the biomass and diversity of weeds. It is therefore important that practitioners understand such variation and apply weed management practices accordingly. Key words: Conventional tillage, reduced tillage, weed composition, weed dynamics.

Data on the population size and trends of large carnivores remains the cornerstone of effective management and conservation programs. However, such data are rarely available for the majority of large carnivore species. Furthermore, large carnivore research is often directed towards formally protected areas. There is therefore a need to improve our knowledge regarding the population ecology of large carnivores in non-protected areas. In this study we use camera trapping in conjunction with spatially explicit mark—recapture models to estimate leopard Panthera pardus density across different land use types in the Waterberg Biosphere, South Africa. Estimated densities (mean ± SE) ranged from 6.59 (± 5.2/100 km2) on a matrix of commercial game and livestock farms to 5.35 (± 2.93/100 km2) and 4.56 (± 1.35/100 km2) on two protected areas (Lapalala and Welgevonden respectively). Although density estimates had large confidence intervals we suggest that these results indicate similar densities across the three site...

Management zones feature prominently in conservation planning, particularly at large spatial scales, but prioritization of areas of concern is required to focus efforts and limited resources. Human-mediated mortality constitutes a major threat to species persistence, particularly for widespread carnivores that undergo harvest and population control, such as the leopard (Panthera pardus). In this study, we evaluated the extent and spatial distribution of legal anthropogenic offtake of leopards to identify de facto refugia and ecological traps across Limpopo Province, South Africa. We defined refugia as management units with offtake levels below an established sustainable harvest rate, and ecological traps as management units with offtake exceeding the sustainable harvest rate. We assessed offtake at three geographical scales using trophy hunting permit records alone, and then in combination with problem leopard permit records to investigate the compounding effect of additional forms of offtake and the potential for management scale mismatching. Across Limpopo Province, high leopard offtake created fewer areas of refuge than ecological traps. Refugia were smaller in size and within close proximity of ecological traps. Human-mediated leopard mortality occurred mostly in prime leopard habitat. Finer-scaled management units resulted in fewer ecological traps and more refugia, and enables authorities to focus conservation attention in areas of concern. Human-mediated leopard mortality exceeded the annual offtake rate considered sustainable. Our study highlights the importance of assessing both the scale and distribution of the harvest, whilst also considering alternative forms of offtake, when devising harvest management strategies. Management scale mismatching and high human-mediated leopard mortality is of particular concern in Limpopo Province, as such, we propose an adaptive, science-based regulatory framework aimed at improving leopard harvest strategies.

Relatively little is known about protozoan parasites in African animals. Here we investigated the occurrence of protozoan parasites in mammals from South Africa. Oocysts of protozoan parasites were detected in 13 of 56 (23%) fecal samples using conventional microscopic examination methods. Cryptosporidium spp. and Cystoisospora spp. were detected in eight (14%) and five (9%) samples, respectively. Mixed parasitic infection of Cryptosporidium spp. and Cystoisospora spp. was recorded in banded mongoose ( Mungos mungo). Cryptosporidium spp. was detected for the first time in cheetah ( Acinonyx jubatus), spotted hyena ( Crocuta crocuta), and African polecat ( Ictonyx striatus). Antibodies to Toxoplasma gondii and Neospora caninum were not detected by enzyme-linked immunosorbent assay in any of 32 sera tested. We detected T. gondii by PCR in tissues of five of 243 (2%) animals: domestic dog ( Canis lupus familiaris), gerbil ( Gerbilliscus spp.), greater kudu ( Tragelaphus strepsiceros), honey badger ( Mellivora capensis), and white-tailed mongoose ( Ichneumia albicauda). Our isolation of T. gondii from white-tailed mongoose and honey badger was a unique finding. All tissue samples were negative for N. caninum. The study increases our knowledge on the occurrence of protozoan parasites in populations of wild and domestic animals in South Africa.

Human-dominated landscapes comprise the bulk of the world’s terrestrial surface and Africa is predicted to experience the largest relative increase over the next century. A multi-scale approach is required to identify processes that maintain diversity in these landscapes. Here we identify scales at which animal diversity responds by partitioning regional diversity in a rural African agro-ecosystem between one temporal and four spatial scales. Human land use practices are the main driver of diversity in all seven animal assemblages considered, with medium sized mammals and birds most affected. Even the least affected taxa, bats and non-volant small mammals (rodents), responded with increased abundance in settlements and agricultural sites respectively. Regional turnover was important to invertebrate taxa and their response to human land use was intermediate between that of the vertebrate extremes. Local scale (< 300 m) heterogeneity was the next most important level for all taxa, highlighting the importance of fine scale processes for the maintenance of biodiversity. Identifying the triggers of these changes within the context of functional landscapes would provide the context for the long-term sustainability of these rapidly changing landscapes.

Rodent pests are especially problematic in terms of agriculture and public health since they can inflict considerable economic damage associated with their abundance, diversity, generalist feeding habits and high reproductive rates. To quantify rodent pest impacts and identify trends in rodent pest research impacting on small-holder agriculture in the Afro- Malagasy region we did a systematic review of research outputs from 1910 to 2015, by developing an a priori defined set of criteria to allow for replication of the review process. We followed the Preferred Reporting Items for Systematic Reviews and Meta-Analyses guidelines. We reviewed 162 publications, and while rodent pest research was spatially distributed across Africa (32 countries, including Madagascar), there was a disparity in number of studies per country with research biased towards four countries (Tanzania [25%], Nigeria [9%], Ethiopia [9%], Kenya [8%]) accounting for 51% of all rodent pest research in the Afro- Malagasy region. There was a disparity in the research themes addressed by Tanzanian publications compared to publications from the rest of the Afro-Malagasy region where research in Tanzania had a much more applied focus (50%) compared to a more basic research approach (92%) in the rest of the Afro-Malagasy region. We found that pest rodents have a significant negative effect on the Afro-Malagasy small-holder farming communities. Crop losses varied between cropping stages, storage and crops and the highest losses occurred during early cropping stages (46% median loss during seedling stage) and the mature stage (15% median loss). There was a scarcity of studies investigating the effectiveness of various management actions on rodent pest damage and population abundance. Our analysis highlights that there are inadequate empirical studies focused on developing sustainable control methods for rodent pests and rodent pests in the Africa-Malagasy context is generally ignored as a research topic.

L.H.S was supported by NRF doctoral (No. 74819) and postdoctoral grants (No. 88179), and L.T. by research funds available at the Vrije University Amsterdam.

There are considerable challenges in the conservation of large carnivores, caused by large area requirements, low reproduction rates and low population densities coupled with their tendency to cause conflict with humans. Trophy hunting is one strategy to increase support for large carnivore conservation. Leopards, Panthera pardus, rank among the most sought-after trophies in South Africa. However, trophy hunting has been suggested as partly responsible for leopard population declines, and leopards are also killed in retaliatory actions. In this study we used a stochastic population model to evaluate the relative influences of retaliatory killing and trophy harvest on leopard population persistence, and to assess the sustainability of the current leopard trophy harvest in South Africa. There was a stronger effect of variation in retaliatory killing than of harvest on population persistence. Although we found low extinction risks for South African leopards within 25 years, high risks of population declines ...

Estimation of survival rates is important for developing and evaluating conservation options for large carnivores. However, telemetry studies for large carnivores are often characterized by small sample sizes that limit meaningful conclusions. We used data from 10 published and 8 unpublished studies of leopards Panthera pardus in southern Africa to estimate survival rates and investigate causes of leopard mortality. Mean survival rates were significantly lower in non-protected (0.55 ± SE 0.08) compared to protected areas (0.88 ± 0.03). Inside protected areas juveniles had significantly lower survival (0.39 ± 0.10) compared to subadults (0.86 ± 0.07) and adults (0.88 ± 0.04). There was a greater difference in cause of death between protected and non-protected areas for females compared to males, with people being the dominant cause of mortality outside protected areas for both females and males. We suggest there is cause for concern regarding the sustainability of leopard populations in South Africa, as high female mortality may have severe demographic effects and a large proportion of suitable leopard habitat lies in non-protected areas. However, because a large proportion of deaths outside protected areas were attributed to deliberate killing by people, we suggest that management interventions may have the potential to increase leopard survival dramatically. We therefore stress the urgency to initiate actions, such as conflict mitigation programmes, to increase leopard survival in non-protected areas.

Carnivores that persist outside of protected areas are often deemed to have highly adaptable and generalist foraging strategies. Using data from three GPS-collared female leopards (Panthera pardus) and over an eight-month period, we attempted to establish whether leopards in a mountainous landscape display preferential behaviour (i.e. specialist) or not (i.e. generalist). We investigated whether female leopards used habitats in accordance with availability for both hunting and regular activities, and whether female leopards demonstrated preference for a particular prey species. Finally, we assessed tree caching behaviour and discuss leopard spatial dynamics in the context of superior competitors, such as lions (Panthera leo). Female leopards demonstrated selection for certain prey species. Mixed closed woodland (greatest vegetation density; lowest prey density) and tall open woodland (greatest vegetation biomass; high prey density) were preferred over other habitats. Female leopards on Welgevonden cached significantly larger prey species in trees. Our results suggest that female leopards in this landscape are selecting habitats - enabling the exploitation of certain ecological features - that balance energy expenditure against the likelihood of capturing prey in an attempt to maximize efficiency, whilst reducing inter-specific competition. We suggest that female leopards are limited in their foraging ability as a result of interference competition by superior species like lions, which in turn, affects their choice of prey. These findings highlight the potential importance of numerous ecological, spatio-temporal, and anthropogenic factors that influence leopard behaviour, and therefore have significant implications for leopard persistence and conservation.

Foraging data for terrestrial carnivorans are most often obtained by the analysis of faecal samples - an established technique. However, advances in satellite tracking technology are allowing researchers to locate kill sites by investigating global positioning system (GPS) clusters in both space and time. Here we compare leopard Panthera pardus dietary estimates (composition and biomass) of small, medium and large prey using three techniques: faecal analysis, GPS cluster analysis and GPS cluster analysis supplemented with faecal samples located at cluster sites. We demonstrate that estimates of leopard prey composition and biomass intake from each technique produce comparatively similar results. Nevertheless, the detection of feeding events did increase by 20-23% when supplementing GPS-located kills with faecal samples. The investigation of GPS clusters offers an equivalent method of leopard dietary estimation to that of faecal analysis. When carried out intensively, the GPS cluster method is capable of detecting leopard predation on prey species within small, medium and large weight categories. Although requiring additional resources, supplementing GPS-located kills with GPS-located faecal samples results in the most detailed dietary estimates by detecting kills missed during GPS cluster investigations.

African wild dog (Lycaon pictus) populations and their geographic distributions have been greatly reduced due to direct human persecution and habitat reduction; however, remnant groups still manage to persist on private reserves and farmland. Farmland, especially game farming areas, can potentially be suitable for wild dog conservation. However, little is known about them in these areas. To increase our understanding regarding the presence and general ecology of wild dogs on private land we used a camera trapping picture database initially used to census leopards (Panthera pardus) to report on wild dog presence on private farmland and reserves within the Waterberg Biosphere, South Africa. We identified nine different wild dogs from the camera trap images and all wild dog signs were restricted to Lapalala. We further supplemented the data set with opportunistically collected scats to report on wild dog prey use. Only three species could be identified in the scats, namely bushbuck (Tragelaphus scriptus), im...

Leopards do not preferentially favour baboons as prey, but they are considered the primary predators of baboons across Africa. Even in areas where baboons are abundant, their contribution to leopard diet seldom exceeds 5% of biomass. It is suggested that the extreme aggressiveness of baboons, group vigilance and their high mobility when escaping may limit leopard predation. Male baboons are particularly aggressive, and retaliation often leads to the death of the leopard. However, evidence suggests that leopards may learn to catch and kill certain dangerous prey. This study reports predation on chacma baboons by 3 female leopards on a private game reserve in the Waterberg Mountains of South Africa. Potential leopard feeding sites were identified using global positioning system (GPS) location clusters obtained from GPS collars. Over a 5-month period, we investigated 200 potential leopard feeding sites and located 96 leopard feeding/kill sites. Baboons constituted 18.7% of the leopards’ biomass intake. The majority of baboons preyed upon were adults and 70% of the kills were diurnal. In terms of the measured variables, there were no significant differences in the way the leopards preyed upon baboons, compared to the rest of the prey species.

The ability to determine the prey-specific biomass intake of large predators is fundamental to their conservation. In the absence of actual prey data, researchers generally use a “unit mass” method (estimated as 3/4 adult female mass) to calculate the biomass intake of predators. However, differences in prey preference and range across geographic regions are likely to have an influence on biomass calculations. Here we investigate the influence of estimated prey mass on leopard biomass calculations, and subsequent carrying capacity estimates, in an understudied mountain population. Potential leopard feeding sites were identified using global positioning system (GPS) location clusters obtained from GPS collars. We investigated 200 potential leopard feeding sites, of which 96 were actual feeding sites. Jaw bones, horns, hooves, and other indicative bones were used to determine gender and age of prey items, which were subsequently used to calculate mass of each prey item based on previously published values. There were significant differences in the biomass values calculated using the traditional unit mass method and the calculated prey masses obtained from leopard feeding sites. However, there were no considerable differences in the carrying capacity estimates using the preferred prey species model and leopard density estimates calculated using a non-biased spatial approach, which suggests that estimating carnivore carrying capacity based on 3/4 adult female masses is a reliable method also for the mountain population in this study.

Worldwide rodent pests are of significant economic and health importance. Controlling rodent pests will, therefore, not only benefit food security but also human and animal health. While rodent pests are most often chemically controlled, there is increased interest in biological control through avian predation. A rich body of research has addressed the impact of avian predators on wild rodent populations, but little is known about the effectiveness of avian predators as biological control agents of rodent pests in agricultural systems. In this study, we systematically reviewed research that investigated different aspects of avian predation on rodent pest populations in order to increase our understanding of the impact and effectiveness of avian predation on rodent pests. Several avian predators (Tyto alba, Elanus axillaris, Falco tinnunculus, Falco cenchroides, Bubo bengalensis, Buteo rufinus) were commonly cited in the biological control of rodents; however, barn owls (T. alba) are the most cited species (86% of studies). We found some support that the use of avian predators produced positive, measurable effects where increased presence of avian predators tended to lower rodent pest numbers, resulting in lower crop damage. However, our review highlighted several shortcomings related to research on avian predation of rodent pests. First, research concerning rodent pest control through avian predation was limited (1.86 articles per year). Secondly, we found that studies lack statistical rigor to detect and measure change in rodent pest species abundance. Finally, the majority of studies were short term and therefore not able to evaluate long term sustainable rodent pest population suppression. We suggest that current shortcomings could be adequately addressed with control-treatment studies that quantitatively investigate the effects of avian predation on rodent pest populations and agricultural impact. Such research could help develop recommendations regarding the use of avian predators in rodent pest management.

Large mammalian carnivores are threatened by anthropogenic environmental impacts, particularly through habitat loss which often cause population declines. Understanding the extent of suitable habitat is therefore of great importance for carnivore conservation. The leopard (Panthera pardus) is a widespread and relatively common large carnivore, but the species is declining in large parts of its range. Using maximum entropy-based habitat models, we estimated the extent of suitable leopard habitat in South Africa, what variables that are associated with suitable leopard habitats, the extent of habitat that has been negatively impacted by human activity and the effectiveness of protected areas to capture suitable habitat. Suitable leopard habitat was highly fragmented. Although vegetation and physical variables were the most influential variables for habitat suitability, livestock farming primarily seem to underlie fragmentation. We suggest that the sustainability of the South African leopard population depends on maintaining dispersal routes between areas with suitable habitat. This will require mitigation of human‐carnivore conflict in habitat corridors, particularly mitigation strategies targeting conflict between carnivores and livestock farmers. Because most suitable habitat occurred outside of protected areas, we also recommend that leopard conservation efforts should focus on areas that are not legally protected.

Apex predators are essential for the viability of healthy ecosystems. By studying carnivoran feeding ecology, we can obtain a better understanding of the ecological limits, resilience and predator–prey dynamics that govern these populations. However, monitoring elusive predators – like the leopard Panthera pardus – is often fraught with logistical and financial constraints, particularly in inaccessible terrain. In this study, we identified clusters of Global Positioning System (GPS) points from four GPS-collared leopards and investigated them in the field for potential kills. Environmental data from cluster sites were gathered alongside spatial and temporal data collected via GPS cluster analysis to develop statistical models capable of predicting the occurrence of leopard predatory events. Our results demonstrate that leopard predation can be accurately modelled either by using a combination of field data (i.e. collected at cluster sites) and remote data (i.e. obtained via GPS analysis), or simply remote data alone. Kills were more likely to be present at clusters where leopards exhibited longer handling times, at sites with dense vegetation cover, when leopards were more active 12 h before the cluster than 12 h after, where more tree refugia were present, in areas of higher elevation, at sites containing low levels of shrub cover, and when clusters began during diurnal or crepuscular hours. By using this modelling approach, comprehensive predation studies are not only efficient and cost-effective but also logistically feasible across a range of different habitats and may even be applied to other carnivorans. It provides a detailed analytical means of studying broader aspects of carnivoran feeding ecology, such as predation habits, carrying capacity, ecological hunting requirements and species interactions, which are important aspects of carnivoran management.