Natal dispersal enables population connectivity, gene flow and metapopulation dynamics. In polygynous mammals, dispersal is typically male-biased. Classically, the ‘mate competition’, ‘resource competition’ and ‘resident fitness’ hypotheses predict density-dependent dispersal patterns, while the ‘inbreeding avoidance’ hypothesis posits density-independent dispersal. In a leopard (Panthera pardus) population recovering from over-harvest, we investigated the effect of sex, population density and prey biomass, on age of natal dispersal, distance dispersed, probability of emigration and dispersal success. Over an 11-year period, we tracked 35 subadult leopards using VHF and GPS telemetry. Subadult leopards initiated dispersal at 13.6 ± 0.4 months. Age at commencement of dispersal was positively density-dependent. Although males (11.0 ± 2.5 km) generally dispersed further than females (2.7 ± 0.4 km), some males exhibited opportunistic philopatry when the population was below capacity. All 13 females were philopatric, while 12 of 22 males emigrated. Male dispersal distance and emigration probability followed a quadratic relationship with population density, whereas female dispersal distance was inversely density-dependent. Eight of 12 known-fate females and 5 of 12 known-fate male leopards were successful in settling. Dispersal success did not vary with population density, prey biomass, and for males, neither between dispersal strategies (philopatry vs. emigration). Females formed matrilineal kin clusters, supporting the resident fitness hypothesis. Conversely, mate competition appeared the main driver for male leopard dispersal. We demonstrate that dispersal patterns changed over time, i.e. as the leopard population density increased. We conclude that conservation interventions that facilitated local demographic recovery in the study area also restored dispersal patterns disrupted by unsustainable harvesting, and that this indirectly improved connectivity among leopard populations over a larger landscape.

Natal dispersal promotes inter-population linkage, and is key to spatial distribution of populations. Degradation of suitable landscape structures beyond the specific threshold of an individual's ability to disperse can therefore lead to disruption of functional landscape connectivity and impact metapopulation function. Because it ignores behavioral responses of individuals, structural connectivity is easier to assess than functional connectivity and is often used as a surrogate for landscape connectivity modeling. However using structural resource selection models as surrogate for modeling functional connectivity through dispersal could be erroneous. We tested how well a second-order resource selection function (RSF) models (structural connectivity), based on GPS telemetry data from resident adult leopard (Panthera pardus L.), could predict subadult habitat use during dispersal (functional connectivity). We created eight non-exclusive subsets of the subadult data based on differing definitions of dispersal to assess the predictive ability of our adult-based RSF model extrapolated over a broader landscape. Dispersing leopards used habitats in accordance with adult selection patterns, regardless of the definition of dispersal considered. We demonstrate that, for a wide-ranging apex carnivore, functional connectivity through natal dispersal corresponds to structural connectivity as modeled by a second-order RSF. Mapping of the adult-based habitat classes provides direct visualization of the potential linkages between populations, without the need to model paths between a priori starting and destination points. The use of such landscape scale RSFs may provide insight into predicting suitable dispersal habitat peninsulas in human-dominated landscapes where mitigation of human-wildlife conflict should be focused. We recommend the use of second-order RSFs for landscape conservation planning and propose a similar approach to the conservation of other wide-ranging large carnivore species where landscape-scale resource selection data already exist.

Polygynous species follow sex-specific spacing patterns to maximize reproductive success, and changes in population density under otherwise stable environmental conditions likely provoke sex-specific responses in spacing patterns. A classical dual reproductive strategy hypothesis posits that female home range size and overlap are set by habitat productivity and remain stable under increasing population density, whereas male home range size and overlap decrease with increased mate competition. An alternative dispersal-regulated strategy predicts that females relinquish part of their home range to philopatric daughters and form matrilineal clusters, while adult male spacing is stable with density-dependent subadult male emigration rates. We used 11 years of telemetry data to assess the response of adult leopard Panthera pardus spacing following the release of harvest pressure. Female annual home ranges and core areas were smaller than in males. Intersexual overlap was larger than intra-sexual overlap in males or in females. As leopard density increased, female home range size and inter-annual fidelity in home range use decreased, and females formed matrilineal kin clusters. In contrast, male leopards maintained large home ranges, and did not track female home range contraction. Spacing dynamics in adult leopards was consistent with dispersal-regulated strategies, and did not support a classical dual reproductive strategy. Our study suggests possible hidden lag effects of harvest disturbance on spacing dynamics that are not necessarily apparent when only assessing demographic recovery of harvested populations.

The risk of transmission of pathogens from free-ranging wild boars (Sus scrofa scrofa) to outdoor domestic pigs (S. scrofa domesticus) is of increasing concern in many European countries. We assess this risk, using Switzerland as an example. We estimated 1) the prevalence of important pathogens in wild boars and 2) the risk of interactions between wild boars and outdoor pigs. First, we tested 252 wild boars from selected areas between 2008 and 2010 for infection with Brucella spp. Bacterial prevalence was estimated to 28.8% (confidence interval [CI] 23.0-34.0) when using bacterial culture (B. suis Biovar 2) and real-time polymerase chain reaction. Antibody prevalence was 35.8% (CI 30.0-42.0), which was significantly higher than in previous studies in Switzerland. We also tested 233 wild boars for porcine reproductive and respiratory syndrome virus (PRRSV). Antibody prevalence was 0.43% (CI 0.01-2.4) for EU-PRRSV and real-time reverse transcription polymerase chain reaction results were negative. These findings suggest that B. suis is increasingly widespread in wild boars and PRRSV is currently not of concern. Second, we documented the spatial overlap between free-ranging wild boars and outdoor piggeries by mapping data on their respective occurrence. Wild boars are most widespread in the mountain range along the western and northern Swiss borders, while most piggeries are located in central lowlands. A risk of interaction is mainly expected at the junction between these two bioregions. This risk may increase if wild boars expand eastward and southward beyond anthropogenic barriers believed to limit their range. Therefore, we evaluated the potential of expansion of the wild boar population. Population trends suggest a continuous increase of wild boars for the past 15 yr. Surveillance of selected wildlife passages using cameras on highways and main roads indicates that these barriers are permeable (average of up to 13 wild boar crossings per 100 days). Thus an increase of wild boar range should be considered. There may be a risk of B. suis spillover from wild boars in Switzerland, which could increase in the future. Data on the occurrence of interactions between pigs and wild boars are needed to assess this risk.

Anthropogenic mortality of wildlife is typically inferred from measures of the absolute decline in population numbers. However, increasing evidence suggests that indirect demographic effects including changes to the age, sex, and social structure of populations, as well as the behavior of survivors, can profoundly impact population health and viability. Specifically, anthropogenic mortality of wildlife (especially when unsustainable) and fragmentation of the spatial distribution of individuals (homeranges) could disrupt natal dispersal mechanisms, with long-term consequences to genetic structure, by compromising outbreeding behavior and gene flow. We investigate this threat in African leopards (Panthera pardus pardus), a polygynous felid with male-biased natal dispersal. Using a combination of spatial (home-range) and genetic (21 polymorphic microsatellites) data from 142 adult leopards, we contrast the structure of two South African populations with markedly different histories of anthropogenically linked mortality. Home-range overlap, parentage assignment, and spatio-genetic autocorrelation together show that historical exploitation of leopards in a recovering protected area has disrupted and reduced subadult male dispersal, thereby facilitating opportunistic male natal philopatry, with sons establishing territories closer to their mothers and sisters. The resultant kin-clustering in males of this historically exploited population is comparable to that of females in a well-protected reserve and has ultimately led to localized inbreeding. Our findings demonstrate novel evidence directly linking unsustainable anthropogenic mortality to inbreeding through disrupted dispersal in a large, solitary felid and expose the genetic consequences underlying this behavioral change. We therefore emphasize the importance of managing and mitigating the effects of unsustainable exploitation on local populations and increasing habitat fragmentation between contiguous protected areas by promoting in situ recovery and providing corridors of suitable habitat that maintain genetic connectivity. K E Y W O R D S home-range, kin-clustering, microsatellites, Panthera pardus, philopatry, relatedness S U PP O RTI N G I N FO R M ATI O N Additional supporting information may be found online in the Supporting Information section. How to cite this article: Naude VN, Balme GA, O'Riain J, et al. Unsustainable anthropogenic mortality disrupts natal dispersal and promotes inbreeding in leopards.

Summary Understanding resource selection and quantifying habitat connectivity are fundamental to conservation planning for both land‐use and species management plans. However, data sets 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 data sets 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 data sets. 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 data sets (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.

Camera traps are a key tool in ecological studies, and are increasingly being used to understand entire communities. However, robust inferences continue to be hampered by low detection of rare and elusive species. Attractants can be used to increase detection rates, but may also alter behaviour, and little research has evaluated short-term, localized response to the presence of attractants. We conducted three camera trap surveys in Kibale National Park, Uganda, using food baits and scent lures (“attractants”) at each camera station to entice small carnivores to pass in front of camera stations. To examine the interrelationship between scavenging and response to attractants, we also placed camera traps at five food refuse pits. We modelled the effect of attractant and duration of trap placement on the detection probability of small carnivores and selected African golden cat Caracal aurata prey items. We examine transient site response of each species, by comparing our observed likelihood of detection in each 24 h period from 1–7 d following refreshing of attractants to randomly generated capture histories. African civet Civettictis civetta , rusty-spotted genet Genetta maculata , African palm civet Nandinia binotata , and marsh mongoose Atilax paludinosus detection probabilities were highest and Weyns’s red duiker Cephalophus wenysi detection probability was lowest immediately after attractants were placed. Within 24 h after attractant was placed, rusty-spotted genet and African palm civet were more likely to be detected and African golden cat, red duiker, and blue duiker Philantomba monticola were less likely to be detected. Our results suggest that attractants can increase detection of small-bodied species and include some arboreal species in terrestrial camera trap sampling. However, attractants may also alter short-term visitation rates of some species, with potentially cascading effects on others. Community level and intraguild interaction studies should control for the potentially confounding effects of attractants on spatial activity patterns.

Density estimates for large carnivores derived from camera surveys often have wide confidence intervals due to low detection rates. Such estimates are of limited value to authorities, which require precise population estimates to inform conservation strategies. Using lures can potentially increase detection, improving the precision of estimates. However, by altering the spatio-temporal patterning of individuals across the camera array, lures may violate closure, a fundamental assumption of capture-recapture. Here, we test the effect of scent lures on the precision and veracity of density estimates derived from camera-trap surveys of a protected African leopard population. We undertook two surveys (a ‘control’ and ‘treatment’ survey) on Phinda Game Reserve, South Africa. Survey design remained consistent except a scent lure was applied at camera-trap stations during the treatment survey. Lures did not affect the maximum movement distances (p = 0.96) or temporal activity of female (p = 0.12) or male leopards (p = 0.79), and the assumption of geographic closure was met for both surveys (p >0.05). The numbers of photographic captures were also similar for control and treatment surveys (p = 0.90). Accordingly, density estimates were comparable between surveys (although estimates derived using non-spatial methods (7.28–9.28 leopards/100km2) were considerably higher than estimates from spatially-explicit methods (3.40–3.65 leopards/100km2). The precision of estimates from the control and treatment surveys, were also comparable and this applied to both non-spatial and spatial methods of estimation. Our findings suggest that at least in the context of leopard research in productive habitats, the use of lures is not warranted.