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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.

As landscape-scale conservation models grow in prominence, assessments of how wildlife utilise multiple-use landscapes are required to inform effective conservation and management planning. Such efforts should incorporate multi-species perspectives to maximise value for conservation, and should account for scale to accurately capture species-environment relationships. We show that the random forest machine learning algorithm can be used to model large-scale sign-based data in a multi-scale framework. We used this method to investigate scale-dependent habitat associations for 16 mammal species of high conservation importance across the southern Kavango Zambezi (KAZA) Transfrontier Conservation Area in Botswana and Zimbabwe. Our findings revealed substantial variation in factors shaping habitat use across species, and illustrate that different species often have divergent responses to the same environmental and anthropogenic factors, and differ in the scales at which they respond to them. For all variables across all species, scale optimisation most often selected our largest scale. Precipitation, soil nutrients, and vegetation appeared to be the most important factors determining mammal distributions, likely through their associations with food resources for herbivores and, in turn, prey availability for carnivores. Anthropogenic pressures also had an important influence, with many species selecting against areas with high cattle density. The variety of relationships with human density indicated that species vary in their tolerance of humans. We found a consistent positive relationship with areas under high protection, and negative relationship with unprotected and less-strictly protected areas. Policy implications. Through a novel application of random forest modelling to spoor data from 16 mammal species, this study highlights the importance of adopting a multi-scale, multi-species approach for decision-making processes that depend on understanding wildlife distributions and habitat associations, such as protected area and corridor prioritisation. The findings identify changing rainfall patterns and increasing livestock numbers as emerging trends that may impact wildlife distributions, both within sub-Saharan Africa and on a global scale. Wildlife management authorities should use modelling exercises and adaptive management to ensure that protected area networks remain fit for purpose under anticipated changes in rainfall under climate change, and explore initiatives that promote coexistence of wildlife and livestock.

Many savanna-dependent species in Africa including large herbivores and apex predators are at increasing risk of extinction. Achieving effective management of protected areas (PAs) in Africa where lions live will cost an estimated US$ >1–2 billion/year in new funding. We explore the potential for fire-management-based carbon financing programs to fill this funding gap and benefit degrading savanna ecosystems. We demonstrate how introducing early dry season fire management programs could produce potential carbon revenues (PCRs) from either a single carbon financing method (avoided emissions) or from multiple sequestration methods ranging from US$ 59.6–655.9 million/year (at US$ 5/ton) or US$ 155.0 million/year to US$ 1.7 billion/year (at US$ 13/ton). We highlight variable but significant PCRs for savanna PAs from US$ 1.5–44.4 million/year per PA. We suggest investing in fire management programs to jump-start the United Nations Decade of Ecological Restoration to help restore degraded African savannas and conserve imperiled keystone herbivores and apex predators.

Social influence impacts individuals’ behavior through norms – the perceived appropriateness of behaviors – and social referents, whose behavior is copied by others. Interventions using social influence may help promote conservation-friendly behaviors. We explored how perceptions are influenced by descriptive norms (what people do) and social referents (who does it) in relation to livestock management in Kenya. By presenting participants with vignettes describing different livestock management scenarios, we explored how two norms (‘high’ vs. ‘low’ standards of livestock management) and three social referents (‘neighbor,’ ‘leader,’ ‘father’) influence participants’ perceptions of aspects of livestock management. More senior social referents and ‘high’ standard norms were associated with greater impact on perceived importance of livestock management behaviors. Age also had an effect, with higher perceived importance and greater effect of norm treatment in younger respondents. Encouraging social seniors to model desired behavior may be a possible strategy for conservation behavioral interventions.

Shifting human–wildlife conflict towards coexistence requires a robust understanding of where conflict happens and why. Spatial models of livestock depredation by wild predators commonly identify depredation hotspots in areas where livestock are most abundant (e.g. nearer villages or pasture). This may reflect underlying livestock distribution, rather than imply these areas are inherently risky for livestock. This limits the predictive power of these models and their usefulness for conflict mitigation and wild carnivore conservation. Here, we build spatial models of both cattle depredation (530 attacks mostly by lions and hyenas; 2009-2013), and cattle presence (14 GPS-collared herds; 2010-2012) near Hwange National Park, Zimbabwe. We use Bayes’ theorem to combine the cattle depredation and presence models to quantify risk as the conditional probability of depredation given livestock presence. Our raw depredation models predicted higher depredation rates where cattle presence was more likely (near villages and in more open habitats). By contrast, our risk model predicted higher risk further from human activity and in more dense vegetation (where depredation rates were higher than expected given the low probability of cattle presence). Risk has also increased sharply towards protected areas (core carnivore habitat). Our formulation of risk captures high-risk areas as those where livestock are most accessible (i.e. vulnerable) to predators as opposed to simply where they are most available (as in much previous work). We make recommendations for livestock protection and wild carnivore conservation based on our quantification of risk, such as where to avoid herding livestock and which areas to prioritize for livestock protection. Our approach may be profitably applied to guide safer livestock grazing or herding in other contexts where depredation and livestock movement data are available. We hope that the concepts and methods that we develop here will help advance the future study and mitigation of human–wildlife conflict more generally.

Olfaction is a key sense, enabling animals to locate forage, select mates, navigate their environment, and avoid predation. Wind is an important abiotic factor that modulates the strength of olfactory information detected by animals. In theory, when airflow is unidirectional, an animal can increase odor detection probability and maximize the amount of olfactory information gained by moving crosswind. Given energetic costs inherent to activity and locomotion, behavioral search strategies that optimize the benefit-cost ratio should be advantageous. We tested whether African lions (Panthera leo) modify their movement directionality and distance according to wind speed and direction during hours of darkness when they are most active. We tracked 29 lions in southern Zimbabwe using GPS collars and deployed a weather station to collect detailed abiotic data. We found that when wind speeds increased lions were more likely to move crosswind. We also found that female lions, which tend to hunt more often than males, traveled farther when wind speeds were stronger. The results of our analysis suggest that lions adjust their movement behavior according to wind speed and direction. We inferred that this was a behavioral decision to maximize the amount of olfactory information gained per unit of energy spent. Our findings not only offer one of the first detailed insights on large carnivore anemotaxis (movement direction relative to wind) but also make an important contribution towards understanding the influence of wind on predator ecology in general which remains understudied to date.

Globally three quarters of large terrestrial mammalian predators are in decline and many populations are data deficient, including those of African leopards across much of their range. Here we assess the drivers of decline African leopard populations in 16 camera trap surveys covering a total area of 15 120 km2, across a gradient of anthropogenic impact, management and geography, in protected areas across the Zimbabwean component of the Kavango-Zambezi (KAZA) Transfrontier Conservation Area. Population density was calculated using spatially explicit mark-recapture estimators and Generalised Additive Models (GAM) were used to assess factors affecting population density. Density estimates ranged from 0.7 to 12.2 (mean 2.9 ± 2.7) leopards/100km2. Leopard density was higher in wooded sites and rugged terrain but negatively affected by human factors including human appropriation of net primary productivity (HANPP), trophy hunting risk and bush-meat poaching. High lion densities (>6.0 lions/ 100km2) negatively affected leopard density. Annual rainfall over a gradient of ~ 300mm across survey sites was not influential in predicting population density. Previous assessments of the drivers of declining leopard population density (CITES 1988), asserting that leopard densities can be predicted by annual rainfall and are unaffected by human disturbance in unmodified habitat are not supported by our findings. We recommend that the 1988 assessment, used to manage CITES leopard trophy hunting export quotas since the late 1980s, should be reviewed.

Apex predators play important roles in ecosystem functioning and, where they coexist, intraguild interactions can have profound effects on trophic relationships. Interactions between predators range from intraguild predation and competition to facilitation through scavenging opportunities. Despite the increased availability of fine-scale GPS data, the determinants and outcomes of encounters between apex predators remain understudied. We used simultaneous GPS data from collared spotted hyaenas (Crocuta crocuta) and African lions (Panthera leo) in Hwange National Park, Zimbabwe, to determine the environmental conditions of the encounters between the two species, which species provoked the encounter, and which species dominated the encounter. Our results show that encounters between hyaenas and lions are mostly resource-related (over a carcass or around waterholes). In the wet season, encounters mainly occur at a carcass, with lions being dominant over its access. In the dry season, encounters mainly occur in the absence of a carcass and near waterholes. Movements of hyaenas and lions before, during, and after these dry-season encounters suggest two interference scenarios: a passive interference scenario whereby both predators would be attracted to waterholes but lions would leave a waterhole used by hyaenas because of prey disturbance, and an active interference scenario whereby hyaenas would actively chase lions from waterhole areas, which are prime hunting grounds. This study highlights the seasonal dynamics of predator interactions and illustrates how the relative importance of negative interactions (interference competition during the dry season) and positive interactions (scavenging opportunities during the wet season) shifts over the course of the year.

Negative interactions with humans resulting from livestock predation is a major factor influencing the decline of African lion Panthera leo populations across Africa. Here we investigate lion depredation within two Maasai communities in southern Kenya where people and lions coexist in the absence of any formal protected areas. We explore the factors that increase the frequency and severity of lion attacks on pastoralists and their livestock and assess the effectiveness of livestock guarding to reduce damage. Finally, we examine in which circumstances lion depredation triggers retaliation by people. Over a period of 26 months, lions attacked livestock 29 times, resulting in 41 livestock deaths and 19 injuries. There were also two attacks on people. Lions preferred cattle over the more numerous sheep and goats. Attacks on livestock occurred mostly during the dry season and were not affected by changes in prey density or variation in pastoral settlement that brought livestock into closer proximity with lions. Livestock were guarded during 48.2% of lion attacks. Active guarding at pasture disrupted the majority of lion attacks, resulting in lower mortality rates. Passive guarding in corrals at night also disrupted attacks but did not lead to lower livestock mortality.

Using an experimental approach, we analyzed the self-reported changes in farmers’ attitudes toward the African lion (Panthera leo) and a community-based lion conservation program (i.e., the Long Shields Community Guardians), before and after implementation. This program protects livestock and mitigates lion depredation through non-lethal means, with the aim of promoting human-lion coexistence. Results indicated a significant positive shift in attitudes of lions by farmers in the treatment group. This was also the case for farmers in what was recognized as a second unplanned treatment group, called the intermediate group, who learned about the program through communication within their peer network. This article contributed to the growing literature on conservation evaluation and demonstrates that people’s attitudes are valuable for assessing the impact of conservation programs.

Translocations are used to mitigate human–wildlife conflict, secure population viability of isolated populations and introduce or reintroduce populations in former or new range. With wild species increasingly confined to small patches of habitat embedded in human-dominated landscapes, the use of translocations is likely to increase. The cheetah Acinonyx jubatus is a large carnivore species with a long history of translocations. As for other species, evaluation of the success of cheetah translocations is complicated by a scarcity of published results, especially of failed attempts. Yet, such information is crucial to improve future translocations. A relatively well documented case is the translocation of alleged problem cheetahs into Matusadona National Park, Zimbabwe, in the early 1990s. In this study we used a combination of survey methods to reassess the status of Matusadona's cheetah population and model current occupancy in relation to densities of competing carnivores and altitude. Our findings indicate this cheetah population has effectively been extirpated, highlighting the importance of thorough planning and standardized long-term monitoring of translocated populations for the understanding of the factors that affect translocation success.

Variation in moonlight affects foraging, hunting and vigilance behaviours in many nocturnal species. Here, we explore the effect of moonlight on the movement and hunting behaviour of African lions. Previous studies found bright moonlight is associated with reduced hunting success; however, those studies were largely undertaken in open habitats where predators are easily seen by prey species on moonlit nights. In this study we explored whether moonlight affected hunting behaviour and success in a largely wooded environment. Measures of short-term lion movements (distance moved, displacement, and path tortuosity) derived from GPS telemetry data were used as indicators of movement behaviour. Field observations of belly distension were used to assess recent food intake. Lions had greater belly distension (indicating feeding success) on dark nights. However, this change in feeding success was not reflected by lion movement patterns – there was no evidence that these changed across moon phases. There was no evidence that lions used more covered habitats on brighter nights to facilitate concealment.

The range-wide decline of lions has led to their conservation becoming a top priority. Protection of free-ranging lion populations is dependent on securing space for lions but also on the ability and desire of local communities to coexist with lions. Our investigation takes a comparative and case study approach to explore the individual and societal desire to maintain current lion populations alongside communities in, or surrounding, Zimbabwe's Hwange National Park, Tanzania's Ruaha National Park, and Kenya's southern Maasailand. Using data from attitudinal questionnaire surveys, we compare the desire to maintain current lion populations as well as the prevalence and success of conservation interventions aimed at increasing human-lion coexistence. In Maasailand, 88% of the respondents expressed a desire to see current lion populations maintained, while only 42% of the respondents in Ruaha and only 5% of the respondents in Hwange expressed this desire. More respondents reported predation by lions (lion predation) on livestock in Maasailand than in Hwange; personal benefits from conservation were greatest in Maasailand; and exposure to conservation education was highest in Ruaha. The Hwange findings were confounded by Zimbabwe's political and economic climate. In Ruaha and Maasailand, communal and individual conservation benefits influenced desired changes to lion population. Once variation between sites was controlled for, twinning personal benefits and conservation education together was most likely to increase an individual's desire to see current lion populations maintained.

Context: There has been limited research identifying large-scale functional connectivity of wildlife populations across sub-Saharan Africa, despite the increased focus on transboundary conservation networks. Objectives: This study set out to assess the functional connectivity of a highly mobile predator of conservation concern across the Kavango-Zambezi transboundary conservation area (KAZA) and the northern Central Kalahari Game Reserve (CKGR), covering almost 300,000km2 of Botswana, Namibia, Zambia and Zimbabwe.Methods: We analysed the nuclear diversity of 204 individual lions from across the metapopulation using Bayesian and multivariate statistics to assess population structure and recent migration. A maximum-likelihood method was used to determine average male dispersal distances to determine the potential for functional connectivity across the region.Results: The results are consistent with work identifying the existence of ecotypic differences between wetland and dryland lions, but also indicate hierarchical population structure further dividing the population into four geographic clusters; the Okavango Delta, the Central Kalahari, Kafue National Park, and the Hwange-Chobe complex. Analysis of dispersal distances of males suggests that reconnecting the clusters through conservation intervention should be successful in improving gene flow and connectivity across the region. Conclusions: While trans-boundary conservation areas may currently have limited gene flow and connectivity, there is potential for the restoration of functional connectivity via the natural dispersal of highly mobile species. However, the matrix of habitat through which such dispersing animals must traverse must be conducive to their movement and survival, highlighting the importance of land outside protected areas for the preservation of highly mobile animals such as lions.

International audience; Long-distance vocalisation is a characteristic of African lion (Panthera leo) behaviour and is important for maintaining territorial boundaries as well as locating distant group members. Vocal signalling is, however, a flexible behaviour that involves varying costs and benefits depending on environmental, social and spatial factors. Motivated by previous data collection limitations, we developed a novel approach to investigate the influence of atmospheric conditions and animal home range geography on lion vocal behaviour using acoustic and accelerometer biologgers. To compensate for the short lifetime of the acoustic biologger, we trained a machine learning model to detect lion roars from long term acceleration signals which yielded over 500 nights of data from 7 individual lions. Analysis of detected roar events revealed that vocalisations occurred mainly at night with a peak just before dawn. The relative likelihood of vocalisation was negatively related to wind speed and temperature and positively related to absolute humidity suggesting that lions preferred to roar under conditions that reduce sound attenuation and thereby maximise calling area. Roar occurrence was found to be dependent on an animal's location relative to its home range with lions demonstrating an apparent avoidance for vocalising beyond the home range boundary. Lions were also more likely to roar repetitively while closer to rivers and water points within their home range. This study is the first of its kind and not only improves the understanding of lion vocal behaviour but can inform new approaches for recording animal vocalisations remotely.

In recent years lion bones have been legally traded internationally to Asian markets from captive-bred sources in South Africa. There are also indications of increasing instances of illegal international trade in wild lion bones. The existence of parallel captive and wild supplies of lion bone are a cause of law enforcement concern regarding the potential for the laundering of illegally sourced bones through legal trade, and present a problem for the assessment of the conservation impact of wild lion bone trade due to the difficulty of determining what market-share wild and captive-bred lion bones account for. Captive-bred and wild lion bone are visually indistinguishable and no reliable method currently exists for distinguishing them. We present a preliminary study that explores the use of DART mass spectrometry as a method to differentiate between captive-bred and wild lion bones. We find that DART is able to differentiate between a batch of captive-bred South African lion bone and a batch of wild lion bone and suggest that DART mass spectrometry shows strong potential as a tool for the regulation and investigation of lion bone trade. Further testing is needed to prove the suitability of this technique. Therefore, we suggest that further research focuses on testing the capability of DART to differentiate between contemporary wild and captive-bred lion bone originating from South Africa, and attempts to identify chemical markers in bone that can be used as indicators of captive-bred origin.

Conservation scientists are increasingly recognizing the need to evaluate the effectiveness of interventions to improve human–wildlife coexistence across different contexts. Here we assessed the long-term efficacy of the Long Shields Community Guardians programme in Zimbabwe. This community-based programme seeks to protect livestock and prevent depredation by lions Panthera leo through non-lethal means, with the ultimate aim of promoting human–lion coexistence. Using a quasi-experimental approach, we measured temporal trends in livestock depredation by lions and the prevalence of retaliatory killing of lions by farmers and wildlife managers. Farmers that were part of the Long Shields programme experienced a significant reduction in livestock loss to lions, and the annual number of lions subject to retaliatory killing by farmers dropped by 41% since the start of the programme in 2013, compared to 2008–2012, before the programme was initiated. Our findings demonstrate the Long Shields programme can be a potential model for limiting livestock depredation by lions. More broadly, our study demonstrates the effectiveness of community-based interventions to engage community members, improve livestock protection and ameliorate levels of retaliatory killing, thereby reducing human–lion conflict.

SUMMARYLions (Panthera leo) in Africa have lost nearly half their population in just the last two decades, and effective management of the protected areas (PAs) where lions live will cost an estimated USD >$1 B/year in new funding. We explore the potential for launching a fire management and habitat restoration carbon-financing program to help fill this PA management funding gap. We demonstrate how introducing early dry season fire management programs could produce potential carbon revenues (PCR) from either a single carbon-financing method (avoided emissions) or from multiple sequestration methods of USD $59.6-$655.9 M/year (at USD $5/ton) or USD $155.0 M–$1.7 B/year (at USD $13/ton). We highlight variable but significant PCR for PAs with the greatest potential for restoring lion numbers between USD $1.5–$44.4 M per PA. We suggest investing in lion-centric fire management programs to jump-start the United Nations Decade of Ecological Restoration and help preserve African lions across their range.SCIENCE FOR SOCIETYThe United Nation’s recently launched the Decade of Ecological Restoration in response to planet-wide land degradation. This study analyses the potential for savanna fire management programs to restore fire regimes that can generate new sources of revenue from carbon financing for chronically under-funded protected areas in Africa with lions, as lions are a key indicator of savanna ecosystem health. We estimated the amount of carbon saved by shifting fires that normally burn in the late dry season (and emit more carbon) to the early dry season (that accrue more carbon in the soil and woody vegetation). Based on current carbon market values we found substantial potential to eliminate or significantly reduce the $>1B annual funding gap needed to save the lion from extinction. Given additional benefits for nature and people from new savanna fire programs, we recommend integrated conservation and development projects direct more funding to some of the least developed countries with high biodiversity and support fire management programs in Africa.

Human behaviour often determines the success of conservation projects, and the emerging discipline of conservation psychology focuses on understanding and influencing this behaviour. Social norms (a group’s perception of the appropriateness of behaviours) are a key influence on human behaviour, and social norms campaigns can often engender population-wide behaviour changes. Human-predator conflict is a major conservation issue, and one in which human behaviour plays a substantial role: high standards of livestock management can considerably lower predation levels. In this paper, we use factor analysis to show that the livestock management normative belief structure of rural livestock owners is highly conserved between populations in Kenya and Zimbabwe. Through cluster analysis, we also show that qualitatively distinct attitudinal groups can be identified, and that some of these groups are common to both regions. Researchers often assume that social landscapes are unique, but we show that this is not the case for livestock management norms. People’s attitudes are also generally assumed to be site-specific, but we found commonalities across different regions, indicating that certain attitude sectors may be present in all livestock owning populations. If livestock management norms and attitude groups are indeed highly conserved between regions, it may be possible to develop standardised tools with which to understand the norms that influence livestock management behaviour, and identify population sectors for targeted interventions. Often, conservation projects have little in-house social science expertise, and social studies are avoided despite the benefits they bring. Here, we demonstrate that standardised approaches may be possible, and could aid the implementation – and success – of conservation interventions.

1. Prey face a trade-off between acquiring food and avoiding predation, but food availability, and therefore its effect, is rarely measured in field studies investigating non-lethal effects of predation. The main aim of this study is to investigate the role of the presence of predators in the functional adjustments of feeding parameters with patch quality in a medium-size herbivore. 2. In Hwange National Park (Zimbabwe), we set up an experiment by manipulating, over two years, patch quality for impala (Aepyceros melampus), a medium-sized herbivore. We assess predation risk by monitoring the presence of three GPS-equipped predators: African lions (Panthera leo), spotted hyaenas (Crocuta crocuta) and African wild dogs (Lycaon pictus). 3. In enriched, fertilised plots the impalas reduced step rates (i.e. the rates of change in feeding stations), and increased their number of bites per feeding station while bite rates were not affected. Thus, the main adjustment of their feeding was the step rate. The total time the impalas spent vigilant appeared to be a good predictor of the variation of their bite rate. Although vigilance caused a reduction in bite rate when at a feeding station, the impalas reduced the relative costs of vigilance by continuing chewing and processing their food when scanning for predators. 4. When predators were in the vicinity, the impalas increased their exclusive vigilance (high-cost vigilance) but not their vigilance while chewing (low-cost vigilance) and decreased their bite rate while their step rate and the number of bites per feeding station did not change significantly. The impalas were thus visually disconnected from their patch, and reduced their bite rate when actually foraging. Exclusive vigilance increased when both lions and hyaenas were in the vicinity, and when wild dogs were nearby. 5. Patterns of vigilance that altered bite rate were linked to the presence of predators during the previous 24h. Over the long term patch quality was the main determinant of the feeding parameters (step rate and bite rate). This study shows how predators, by affecting the time prey devote to predator detection, shape the functional adjustments of food acquisition by prey to local patch quality.

Conflict with humans is one of the major threats facing the world's remaining large carnivore populations, and understanding human attitudes is key to improving coexistence. We surveyed people living near Hwange National Park about their attitudes toward coexisting with lions. We used ordinal regression models with the results of the survey to investigate the importance of a range of tangible and intangible factors on attitudes. The variables investigated included the costs and benefits of wildlife presence, emotion, culture, religion, vulnerability, risk perception, notions of responsibility, and personal value orientations. This was for the purpose of effectively tailoring conservation efforts but also for ethical policy making. Intangible factors (e.g., fear and ecocentric values) were as important as, if not more important than, tangible factors (such as livestock losses) for understanding attitudes, based on the effect sizes of these variables. The degree to which participants' fear of lions interfered with their daily activities was the most influential variable. The degree to which benefits accrue to households from the nearby protected area was also highly influential, as was number of livestock lost, number of dependents, ecocentric value orientation, and participation in conflict mitigation programs. Contrary to what is often assumed, metrics of livestock loss did not dominate attitudes to coexistence with lions. Furthermore, we found that socioeconomic variables may appear important when studied in isolation, but their effect may disappear when controlling for variables related to beliefs, perceptions, and past experiences. This raises questions about the widespread reliance on socioeconomic variables in the field of human-wildlife conflict and coexistence. To facilitate coexistence with large carnivores, we recommend measures that reduce fear (through education and through protective measures that reduce the need to be fearful), reduction of livestock losses, and ensuring local communities benefit from conservation. Ecocentric values also emerged as influential, highlighting the need to develop conservation initiatives tailored to local values.Importancia de los Factores Tangibles e Intangibles en la Coexistencia Humanos-Carnívoros Resumen Los conflictos con humanos son una de las principales amenazas que enfrentan las poblaciones permanentes de grandes carnívoros del planeta y el entendimiento de las actitudes humanas es importante para mejorar la coexistencia. Preguntamos a las personas que viven cerca del Parque Nacional Hwange sobre su postura hacia la coexistencia con leones. Usamos modelos de regresión ordinal en los resultados del cuestionario para investigar la importancia de una gama de factores tangibles e intangibles para estas posturas. Las variables investigadas incluyeron el costo y beneficio de la presencia de fauna, emociones, cultura, religión, vulnerabilidad, percepción de riesgos, nociones de la responsabilidad y orientaciones de valor personal. Realizamos esto con el propósito de diseñar efectivamente los esfuerzos de conservación pero también para la generación de políticas éticas. Los factores intangibles (p. ej.: el miedo y los valores ecocéntricos) fueron tan importantes, si no es que más importantes, como los factores tangibles (como la pérdida de cabezas de ganado) para el entendimiento de las actitudes, con base en los tamaños del efecto de estas variables. El grado al que el miedo que los participantes sienten por los leones interfiere con sus actividades diarias fue la variable con mayor influencia. El grado al que los hogares del área protegida cercana devengan beneficios también tuvo una influencia alta, como lo tuvo el número de cabezas de ganado perdidas, el número de dependientes, la orientación de los valores ecocéntricos y la participación en programas de mitigación del conflicto. Contrario a lo que frecuentemente se asume, las medidas de la pérdida de cabezas de ganado no dominó sobre las actitudes por la coexistencia con leones. Más allá, encontramos que las variables socioeconómicas pueden parecer importantes cuando se estudian de manera aislada pero su efecto puede desaparecer cuando se controlan las variables relacionadas con las creencias, percepciones y experiencias pasadas. Esto genera preguntas sobre la dependencia hacia las variables socioeconómicas en el área de estudios de conflictos y coexistencia entre humanos y fauna. Para facilitar la coexistencia con carnívoros mayores recomendamos tomar medidas que reduzcan el miedo (por medio de la educación y medidas de protección que reduzcan la necesidad de vivir con miedo), la disminución de la pérdida de cabezas de ganado y garantizarles a las comunidades locales los beneficios que proporciona la conservación. Los valores ecocéntricos también surgieron como influyentes, lo que resalta la necesidad de desarrollar iniciativas de conservación adaptadas a los valores locales.

Trade-driven killing for body parts has long been a major cause of population decline for a number of big cat species. There are now worrying suggestions that commercialised illegal trade in body parts might become a threat for wild lions in Africa, and recent concerns have been raised that trade in captive-bred lion skeletons from South Africa may have stimulated demand for lion bones, claws, and teeth and thus incentivised commercially-driven, targeted poaching of wild lions for illegal trade. However, analysis of the prevalence of commercially-driven, targeted killing of lions for body parts is currently lacking for most major lion populations. In this study we make use of detailed, long-term records of mortality and body part removal from field sites in two of Africa’s lion population strongholds, the Hwange National Park and surrounds, Zimbabwe, within the Kavango Zambezi (KAZA) Transfrontier Conservation Area and the Ruaha landscape in Tanzania, to evaluate potential cases of commercially-driven part removal and targeted killings. We find no evidence of systemic targeted killing of lions for commercial trade at either site and suggest that the majority of part removals are opportunistic and culturally-driven. Nevertheless, we stress the requirement for vigilance around the issue of lion trade and support the development of further range-wide capacity to record wild lion mortality and body part removal data.

International audience; Numerous unknown factors influence anthrax epidemiology in multi-host systems, especially at wildlife/livestock/human interfaces. Serology tests for anti-anthrax antibodies in carnivores are useful tools in identifying the presence or absence ofBacillus anthracisin a range. These were employed to ascertain whether the disease pattern followed the recognized high- and low-risk anthrax zonation in Zimbabwe and also to establish whether anthrax was absent from Hwange National Park in which there have been no reported outbreaks. African lions (Panthera leo)(n = 114) drawn from free-range protected areas and captive game parks located in recognized high- and low-risk zones across Zimbabwe were tested for antibodies to anthrax PA antigen using the ELISA immunoassay. A random selection of 27 lion sera samples comprising 17 seropositive and 10 seronegative sera was further tested in the species-independent toxin neutralization assay (TNA) in order to validate the former as a surveillance tool for anthrax in African lions. Using the ELISA-PA immunoassay, 21.9% (25/114) of the lions tested positive for antibodies to anthrax. Seropositivity was recorded in all study areas, and there was no significant difference (p = .852) in seropositivity between lions in high- and low-risk anthrax zones. Also, there was no significant difference (McNemar's chi-square test = 0.9,p = .343) in the proportion of lions testing positive to anti-PA anthrax antibodies on ELISA-PA immunoassay compared with the TNA, with fair agreement between the two tests [kappa (K) statistic = 0.30; 0.08 < K

A controversial, multifaceted debate surrounds the trade in commercially captive-bred (farmed) lion skeletons. A prominent topic relates to relative preferences for tiger and lion bone in Asian consumer countries. To contribute preliminary information on this subject we conducted the first quantitative study to assess the consumer preferences of the urban public in China and Vietnam for lion versus tiger and wild versus farmed bone wine products. Using an online questionnaire we ranked respondents’ stated preference for wild tiger, farmed tiger, wild lion, and farmed lion bone, and tested for the effect of demographic and attitudinal variables on product preferences. Our findings indicate that in both China and Vietnam tiger bone wine is greatly preferred over lion bone wine, and that respondents showed high levels of fidelity to their choice of farmed or wild designation across species. We emphasise the real-world complexity of lion and tiger bone product interactions and highlight opportunities for further in-depth study.

1. Environmental variations can influence the extent to which individuals interact with other individuals by changing the value of grouping. It is well known that many species can form and disband groups, often in response to the distribution and abundance of resources.2. While previous studies showed that resources influence the broad‐scale structure of animal groups, knowledge gaps remain on whether they affect fine‐scale patterns of association among individuals within groups.3. We quantify association patterns in African lions while simultaneously monitoring the abundance and distribution of prey. We test how social and ecological factors, including individual trait (age, sex, reproductive state) similarity and prey availability (prey abundance, dispersion, herd size and body size) affect within‐pride social structure in African lions.4. We found that individual decisions about associates depended on resource availability with individuals associating equally across all members of the pride when prey herds were scarce, aggregated or large bodied, and associating more exclusively (in subgroups of preferred associates) when prey herds were abundant, dispersed or small bodied. Individuals within lion prides seemed to be buffering against changes in prey availability by modulating their strength and density of connections with conspecifics when prides split into subgroups. The strength and density of connections among individuals within subgroups was greater when prey herds were large and lower when prey herds were dispersed or are large bodied.5. Our findings suggest that individual lions are making social decisions at both the subgroup level and the pride level, with decisions representing putatively fitness‐enhancing strategies. Individuals were typically shifting between having few strong connections and having many weaker connections depending on prevailing ecological conditions, with prey abundance, dispersion and body size having the greatest impact on decisions about splitting into subgroups. The maintenance of connections within prides and subgroups in the face of ecological change suggests that the fission–fusion nature of lion prides might be essential for the long‐term maintenance of social connections even when short‐term conditions do not allow them. More broadly, our study reveals how fission–fusion dynamics and ecological factors can simultaneously have an effect on animals across multiple levels of sociality. published

Transfrontier conservation areas (TFCAs) have the potential to provide havens for large carnivores while preserving connectivity across wider mixed-use landscapes. However, information on the status of species in such landscapes is lacking, despite being a prerequisite for effective conservation planning. We contribute information to this gap for leopard (Panthera pardus) in Africa, where the species is facing severe range contractions, using data from transect surveys of a 30,000km2 area across Botswana and Zimbabwe in the Kavango-Zambezi (KAZA) TFCA. We used occupancy models to assess how biotic, anthropogenic, and management variables influence leopard habitat use, and N-mixture models to identify variables influencing the species' relative abundance. Leopard were detected in 184 out of 413 sampling units of 64km2; accounting for imperfect detection resulted in mean detection probability p ¯ = 0.24 (SD = 0.06) and mean probability of site use ψ ¯ = 0.89 (SD = 0.20). Habitat use was positively influenced by prey availability and high protection. Relative abundance was best predicted by trophy hunting, which had a negative influence, while abundance was positively associated with high protection and availability of steenbok. Our findings suggest that securing prey populations should be a priority in conservation planning for leopard in Africa, and underline the necessity of preserving highly-protected areas within mixed-use landscapes as strongholds for large carnivores. Our findings also support calls for better assessment of leopard population density in trophy hunting areas, and illustrate the value of N-mixture models to identify factors influencing relative abundance of large carnivores.

We explored the perceptions held by subsistence farmers (living in communal lands within the CAMPFIRE programme area in northwestern Zimbabwe) towards the African lion (Panthera leo) and related conservation interventions undertaken by management authorities. Conceptually guided by the cognitive hierarchy, we used a semi-structured, face-to-face interview format to collect data across three different farming communities bordering Hwange and Zambezi National Parks. Ordinal regression models were used to analyse the data. Our results illustrate that farmers' perceptions towards lions were strongly negative and appeared to be associated with the geographic location in which the farmer lived, as well as the farmer's ethnic group. We also found that perceptions towards lions were not associated with specific livestock losses or to the potential benefits farmers received from wildlife conservation, e.g. school classroom blocks or road improvements. Instead, we suggest that fear of lions and perceived risk to livestock or human wellbeing may play a stronger role in shaping farmers' perceptions compared to actual livestock losses. Moreover, we suggest that sharing information across farmer social networks within a community area, along with the potential for media attention over sensational events, may also influence perceptions towards lions. Our results contribute a baseline dataset for future applied research in this area, and provide insight into developing locally-meaningful conservation interventions, including the type of information to be shared, channels for communication, and the benefits derived from participating in wildlife conservation.

Leopard (Panthera pardus) conservation has a strong international dimension. Hunting trophy export quotas established for African range states under the Convention on International Trade in Endangered Species (CITES) are a case in point. We test these quotas, and the methods for their establishment, against the benchmark of the general principles of precaution, sustainable use and adaptive management. The various national approaches and the CITES regime condoning them largely fail this test. For decades, CITES bodies have endorsed apparently arbitrary quotas lacking robust scientific bases, without regular adjustment. Thus, the quotas have been inadequately performing their assigned function within the Convention’s framework. The way in which the CITES leopard quota regime has been operating is fundamentally at odds with the principles of sustainable use, precaution and adaptive management. To remedy this, we offer recommendations on how to embed a science-based, sustainable, precautionary and adaptive approach to quota-setting within the CITES system.

International audience; In large mammal communities, little is known about modification of interspecific interactions through habitat structure changes. We assessed the effects of African elephants (Loxodonta africana) on features of woody habitat structure that can affect predator–prey interactions. We then explored how this can influence where African lions (Panthera leo) kill their prey. Indeed, lions are stalk-and-ambush predators and habitat structure and concealment opportunities are assumed to influence their hunting success. During 2 years, in Hwange National Park, Zimbabwe, kill sites (n = 167) of GPS-collared lions were characterized (visibility distance for large mammals, distance to a potential ambush site and presence of elephant impacts). We compared characteristics of lion kill sites with characteristics of random sites (1) at a large scale (i.e. in areas intensively used by lions, n = 418) and (2) at the microhabitat scale (i.e. in the direct surrounding available habitat

International audience; Previous research has shown that African lions (Panthera leo) have the ability to discriminate between conspecific vocalisations, but little is known about how individual identity is conveyed in the spectral structure of roars. Using acoustic – accelerometer biologgers that allow vocalisations to be reliably associated with individual identity, we test for vocal individuality in the fundamental frequency (f0) of roars from 5 male lions, firstly by comparing simple f0 summary features and secondly by modelling the temporal pattern of the f0 contour. We then assess the application of this method for discriminating between individuals using passive acoustic monitoring. Results indicate that f0 summary features only allow for vocal discrimination with 70.7% accuracy. By comparison, vocal discrimination can be achieved with an accuracy of 91.5% based on individual differences in the temporal pattern of the f0 sequence. We further demonstrate that passively recorded lion roars can be localised and differentiated with similar accuracy. The existence of individually unique f0 contours in lion roars and their relatively lower attenuation indicates a likely mechanism enabling individual lions to identify conspecifics over long distances. These differences can be exploited by researchers to track individuals across the landscape and thereby supplement conventional lion monitoring approaches.

Simple Summary Acting to preserve biodiversity can involve harming individual animals. It has recently been argued that conventional practice has placed too much emphasis on the preservation of collective entities, such as populations and species, at the expense of suffering for individuals. At least some advocates of the ‘Compassionate Conservation’ movement find any deployment of lethal measures in the interests of conservation to be unacceptable. This shifts the balance of priorities too far. While conservationists have a duty to minimise harm, and to use non-lethal measures where feasible, there will be serious implications for conservation if this movement were to be widely influential. Furthermore, the ‘do-no-harm’ maxim the compassionate conservationists advocate does not always promote the welfare of individual animals. Abstract Human activity affecting the welfare of wild vertebrates, widely accepted to be sentient, and therefore deserving of moral concern, is widespread. A variety of motives lead to the killing of individual wild animals. These include to provide food, to protect stock and other human interests, and also for sport. The acceptability of such killing is widely believed to vary with the motive and method. Individual vertebrates are also killed by conservationists. Whether securing conservation goals is an adequate reason for such killing has recently been challenged. Conventional conservation practice has tended to prioritise ecological collectives, such as populations and species, when their interests conflict with those of individuals. Supporters of the ‘Compassionate Conservation’ movement argue both that conservationists have neglected animal welfare when such conflicts arise and that no killing for conservation is justified. We counter that conservationists increasingly seek to adhere to high standards of welfare, and that the extreme position advocated by some supporters of ‘Compassionate Conservation’, rooted in virtue ethics, would, if widely accepted, lead to considerable negative effects for conservation. Conservation practice cannot afford to neglect consequences. Moreover, the do-no-harm maxim does not always lead to better outcomes for animal welfare.

Human–carnivore conflict (HCC) represents one of the greatest threats to rural livelihoods and the persistence of large carnivores. The application of aversive conditioning, the association of unpleasant stimuli with the occurrence of unwanted behaviors, to mitigate HCC has achieved mixed results within and across species, making a better understanding of the factors driving intervention success critical to inform management practices. We explored the degree to which the chasing of African lions (Panthera leo) out of no tolerance zones conditions lion behavior to reduce their rate of return into community lands or rate of repeated livestock killing, providing evidence‐based understanding of program outcomes. We used data from 15 global positioning system (GPS)‐collared lions adjacent to Hwange National Park, Zimbabwe, with each lion receiving 0–17 conditioning treatments, and analyzed the data using recurrent event survival analysis and logistic regression. Chases were most successful (i.e., lion pushed out of the no tolerance zone by sunrise of the following day) in the dry season (i.e., when wild prey were more predictable), in areas closer to the park, and for individuals from smaller and more stable prides (i.e., had not lost a pride male within six months). Adult females and subadult males were more likely than adult males to reenter community lands, and subadult males were most likely to repeatedly depredate livestock. While livestock depredation has decreased since program initiation, the individuals in this study were overall not less likely to enter community lands or depredate livestock in response to chases when chases were considered isolated events. Rather, it was the consistency of deterrence events that proved most important in reducing livestock depredations, likely because of a stronger reinforcement between the undesired behavior and the negative stimulus. However, lions that had previously habitually killed livestock had greater depredation rates even after several conditioning treatments. Aversive conditioning holds promise in the management of carnivores that depredate livestock, but intervention must be consistent, ideally early in the development of problem behaviors, to maximize intervention effectiveness. Methods that separate wildlife from people (i.e., fencing, livestock enclosure fortification), in combination with aversive conditioning, may be needed to provide a sustainable, long‐term solution.

What does trophy hunting (selective hunting for recreation) contribute to wild lion conservation? Macdonald (Report on Lion Conservation with Particular Respect to the Issue of Trophy Hunting. WildCRU, Oxford, UK, 2016) summarises what we know. We identify unknowns, gaps in the knowledge that inhibit conservation planning, including: the causes of lion mortality, the amount of land used for lion trophy hunting, the extent to which trophy hunting depends on lions for financial viability, and the vulnerability of areas used for hunting to conversion to land not used for wildlife, if trophy hunting ceased. The cost of reversing biodiversity loss exceeds income from tourism, including hunting. New financial models are needed, particularly in view of the expanding human population in Africa.

Using surveys of experts associated with 186 sites across 24 countries, we assessed the effectiveness of African protected areas (PAs) at conserving lions and their prey, identified factors that influence conservation effectiveness, and identified patterns in the severity of various threats. Less than one third of sampled PAs conserve lions at ≥ 50% of their estimated carrying capacity (K), and less than half conserve lion prey species at ≥ 50% of K. Given adequate management, PAs could theoretically support up to 4 × the total extant population of wild African lions (~ 83,000), providing a measurable benchmark for future conservation efforts. The performance of PAs shows marked geographic variation, and in several countries there is a need for a significant elevation in conservation effort. Bushmeat poaching was identified as the most serious threat to both lions and to wildlife in general. The severity of threats to wildlife in PAs and the performance of prey populations were best predicted by geographic-socioeconomic variables related to the size of PAs, whether people were settled within PAs, human/livestock densities in neighbouring areas and national economic indicators. However, conservation outcomes for lions were best explained by management variables. PAs tended to be more effective for conserving lions and/or their prey where management budgets were higher, where photographic tourism was the primary land use, and, for prey, where fencing was present. Lions and prey fared less well relative to their estimated potential carrying capacities in poorer countries, where people were settled within PAs and where PAs were used for neither photographic tourism nor trophy hunting.

In a world of shrinking habitats and increasing competition for natural resources, potentially dangerous predators bring the challenges of coexisting with wildlife sharply into focus. Through interdisciplinary collaboration between authors trained in the humanities, social sciences and natural sciences, this paper offers a review of current approaches and a vision for future approaches to understanding and mitigating adverse human-predator encounters. The paper first reviews some limitations to current approaches to mitigation. Second, it reviews an emerging interdisciplinary literature, identifying key perspectives on how to better frame and therefore successfully mitigate such conservation conflicts. Third, it discusses the implications for future research and management practice. It is concluded that a demand for rapid, ‘win-win’ solutions for conservation and development favours dispute resolution and technical fixes, obscuring important underlying drivers of conflicts. Without due cognisance of these underlying drivers, our well intentioned efforts, focussed on ‘human wildlife conflicts,’ will fail.

It has been proposed that there is a thermal cost of the mane to male lions, potentially leading to increased body surface temperatures (T s), increased sperm abnormalities, and to lower food intake during hot summer months. To test whether a mane imposes thermal costs on males, we measured core body temperature (T b) continuously for approximately 1 year in 18 free‐living lions. There was no difference in the 24‐hr maximum T b of males (n = 12) and females (n = 6), and males had a 24‐hr mean T b that was 0.2 ± 0.1°C lower than females after correcting for seasonal effects. Although feeding on a particular day increased 24‐hr mean and 24‐hr maximum T b, this phenomenon was true of both male and female lions, and females had higher 24‐hr mean and 24‐hr maximum T b than males, on both days when lions did not feed, and on days when lions did feed. Twenty‐four‐hour T b was not influenced by mane length or color, and 24‐hr mean T b was negatively correlated with mane length. These data contradict the suggestion that there exists a thermal cost to male lions in possessing a long dark mane, but do not preclude the possibility that males compensate for a mane with increased heat loss. The increased insulation caused by a mane does not necessarily have to impair heat loss by males, which in hot environments is primarily through respiratory evaporative cooling, nor does in necessarily lead to increased heat gain, as lions are nocturnal and seek shade during the day. The mane may even act as a heat shield by increasing insulation. However, dominant male lions frequent water points more than twice as often as females, raising the possibility that male lions are increasing water uptake to facilitate increased evaporative cooling. The question of whether male lions with manes compensate for a thermal cost to the mane remains unresolved, but male lions with access to water do not have higher T b than females or males with smaller manes.