The semi-arid ecosystem of Asiatic Lion Landscape in Saurashtra, Gujarat: Population density, biomass and conservation of nine wild prey species.

The present study aimed to assess the population density, structure, and population change of nine wild prey species in the semi-arid landscape of Saurashtra, Gujarat, India. A total of eight sites, representing a gradient from highly protected woodlands and grasslands to unreserved grasslands, were selected for sampling. We employed the road transect methodology under a distance sampling framework to achieve our objectives. We evaluated the realized growth rate of the Gir ungulate population through linear regression analysis. Our findings revealed that deer species exhibited higher density and biomass in woodlands compared to grasslands and coastal forests. On the other hand, antelopes showed higher density and biomass in grasslands and coastal forests compared to woodlands. The density gradient of wild prey species was influenced by various factors, including habitat structure, social organization, grouping tendencies, and topography. Over the last four decades, the population of wild prey species in Gir showed minimal changes. Our study provides a comprehensive understanding of wild prey species' density and biomass patterns at the landscape level. The inclusion of findings from ecologically significant and unique areas, such as coastal forests, further enhances the importance of this study. The implications of this study extend beyond the conservation of wild prey species alone; they also contribute to the conservation of the large carnivore guild in the Saurashtra landscape.

A pan-African spatial assessment of human conflicts with lions and elephants.

African lions (Panthera leo) and African savanna (Loxodonta africana) and forest (L. cyclotis) elephants pose threats to people, crops, and livestock, and are themselves threatened with extinction. Here, we map these human-wildlife conflicts across Africa. Eighty-two percent of sites containing lions and elephants are adjacent to areas with considerable human pressure. Areas at severe risk of conflict (defined as high densities of humans, crops, and cattle) comprise 9% of the perimeter of these species' ranges and are found in 18 countries hosting, respectively, ~ 74% and 41% of African lion and elephant populations. Although a variety of alternative conflict-mitigation strategies could be deployed, we focus on assessing the potential of high-quality mitigation fences. Our spatial and economic assessments suggest that investments in the construction and maintenance of strategically located mitigation fences would be a cost-effective strategy to support local communities, protect people from dangerous wildlife, and prevent further declines in lion and elephant populations.

A multispecies assessment of wildlife impacts on local community livelihoods.

Conflicts between the interests of agriculture and wildlife conservation are a major threat to biodiversity and human well-being globally. Addressing such conflicts requires a thorough understanding of the impacts associated with living alongside protected wildlife. Despite this, most studies reporting on human-wildlife impacts and the strategies used to mitigate them focus on a single species, thus oversimplifying often complex systems of human-wildlife interactions. We sought to characterize the spatiotemporal patterns of impacts by multiple co-occurring species on agricultural livelihoods in the eastern Okavango Delta Panhandle in northern Botswana through the use of a database of 3264 wildlife-incident reports recorded from 2009 to 2015 by the Department of Wildlife and National Parks. Eight species (African elephants [Loxodonta africana], hippopotamuses [Hippopotamus amphibious], lions [Panthera leo], cheetah [Acinonyx jubatus], African wild dogs [Lycaon pictus], hyenas [Crocuta crocuta], leopards [Panthera pardus], and crocodiles [Crocodylus niloticus]) appeared on incident reports, of which 56.5% were attributed to elephants. Most species were associated with only 1 type of damage (i.e., either crop damage or livestock loss). Carnivores were primarily implicated in incident reports related to livestock loss, particularly toward the end of the dry season (May-October). In contrast, herbivores were associated with crop-loss incidents during the wet season (November-April). Our results illustrate how local communities can face distinct livelihood challenges from different species at different times of the year. Such a multispecies assessment has important implications for the design of conservation interventions aimed at addressing the costs of living with wildlife and thereby mitigation of the underlying conservation conflict. Our spatiotemporal, multispecies approach is widely applicable to other regions where sustainable and long-term solutions to conservation conflicts are needed for local communities and biodiversity.

Article Impact Statement: Impacts of protected wildlife on local community livelihoods can vary and overlap across species, spatial scales, and time of the year. Una Evaluación Multiespecie de los Impactos de la Fauna sobre el Sustento de la Comunidad Local Resumen Los conflictos entre los intereses de la agricultura y la conservación de fauna son una gran amenaza para la biodiversidad y el bienestar humano en todo el mundo. Para tratar estos conflictos se requiere un entendimiento exhaustivo de los impactos asociados con la convivencia con fauna protegida. A pesar de esto, la mayoría de los estudios que reportan sobre los impactos humano-fauna y las estrategias que se usan para mitigarlos se enfocan en una sola especie, lo que simplifica demasiado los complejos sistemas de interacciones humano-fauna. Buscamos caracterizar los patrones espaciotemporales de los impactos por múltiples especies coocurrentes sobre el sustento agrícola en la franja oriental del Delta del Okavango al norte de Botswana mediante el uso de una base de datos de 3,264 reportes de incidentes con fauna registrados entre 2009 y 2015 por el Departamento de Vida Silvestre y Parques Nacionales. Ocho especies (elefante africano [Loxodonta africana], hipopótamo [Hippopotamus amphibious], león [Panthera leo], chita [Acinonyx jubatus], licaón [Lycaon pictus], hiena [Crocuta crocuta], leopardo [Panthera pardus] y cocodrilo [Crocodylus niloticus]) aparecieron en los reportes de incidentes, de los cuales el 56.5% estaba atribuido a los elefantes. La mayoría de las especies estuvo asociada sólo con un tipo de daño (es decir, daño a cultivos o pérdida de ganado). Los carnívoros fueron los principales implicados en los reportes de incidentes relacionados con la pérdida de ganado, particularmente hacia el final de la temporada seca (mayo-octubre). Al contrario, los herbívoros estuvieron asociados con los incidentes de pérdida de cultivos durante la temporada de lluvias (noviembre-abril). Nuestros resultados ejemplifican cómo las comunidades locales pueden enfrentar diferentes dificultades en su sustento por parte de diferentes especies durante diferentes periodos en el año. Tal evaluación multiespecie tiene consecuencias importantes para el diseño de las intervenciones de conservación enfocadas en la resolución de los efectos de la convivencia con la fauna y por lo tanto la mitigación del conflicto de conservación subyacente. Nuestro enfoque multiespecie espaciotemporal puede aplicarse ampliamente a otras regiones en donde las comunidades y la biodiversidad local necesitan soluciones sustentables y a largo plazo para los conflictos de conservación.

Conservation prioritization can resolve the flagship species conundrum.

Conservation strategies based on charismatic flagship species, such as tigers, lions, and elephants, successfully attract funding from individuals and corporate donors. However, critics of this species-focused approach argue it wastes resources and often does not benefit broader biodiversity. If true, then the best way of raising conservation funds excludes the best way of spending it. Here we show that this conundrum can be resolved, and that the flagship species approach does not impede cost-effective conservation. Through a tailored prioritization approach, we identify places containing flagship species while also maximizing global biodiversity representation (based on 19,616 terrestrial and freshwater species). We then compare these results to scenarios that only maximized biodiversity representation, and demonstrate that our flagship-based approach achieves 79-89% of our objective. This provides strong evidence that prudently selected flagships can both raise funds for conservation and help target where these resources are best spent to conserve biodiversity.

Reproductive cycle stage assessment using vaginal cytology evaluation in African lions (Panthera leo).

Vaginal cytology evaluation is an economic, non-invasive technique for indirect monitoring of fluctuations in estrogen concentrations, and thus progression of the estrous cycle. This technique is widely used in domestic dogs for determining timing of artificial insemination. There, however, are only a few reports on the vaginal cytology of non-domestic felids, including lions. This study was conducted, therefore, to describe the vaginal epithelial changes throughout the reproductive cycle of African lions, and to investigate the efficacy of vaginal cytology assessments for predicting reproductive stages. During a 12-month period, reproductive behavioral data and vaginal swabs were collected daily from five lionesses. In total, 541 vaginal smears were evaluated for the proportion of mucosal epithelial cells, neutrophils, bacterial cells, and amount of mucous, cellular debris. One single swab with a large proportion of superficial cells, absence of neutrophils, large number of bacteria, without cellular debris was sufficient for detecting lionesses in estrus. Likewise, one cytology sample with a large proportion of parabasal and intermediate cells, few neutrophils, few bacteria, and large amount of mucous, cellular debris enabled detection of females in advanced diestrus or gestation. To distinguish lionesses in early diestrus from those in an inter-estrous period, at least two consecutive swabs were necessary for satisfactory classification. Overall, evaluation of vaginal cytology samples was an effective technique for differentiation among different stages of the reproductive cycle, confirmation of estrus, and pregnancy diagnosis in lionesses. This technique, therefore, has the potential for application in classifying different stages of the reproductive cycle in other feline species.

Born captive: A survey of the lion breeding, keeping and hunting industries in South Africa.

Commercial captive breeding and trade in body parts of threatened wild carnivores is an issue of significant concern to conservation scientists and policy-makers. Following a 2016 decision by Parties to the Convention on International Trade in Endangered Species of Wild Fauna and Flora, South Africa must establish an annual export quota for lion skeletons from captive sources, such that threats to wild lions are mitigated. As input to the quota-setting process, South Africa's Scientific Authority initiated interdisciplinary collaborative research on the captive lion industry and its potential links to wild lion conservation. A National Captive Lion Survey was conducted as one of the inputs to this research; the survey was launched in August 2017 and completed in May 2018. The structured semi-quantitative questionnaire elicited 117 usable responses, representing a substantial proportion of the industry. The survey results clearly illustrate the impact of a USA suspension on trophy imports from captive-bred South African lions, which affected 82% of respondents and economically destabilised the industry. Respondents are adapting in various ways, with many euthanizing lions and becoming increasingly reliant on income from skeleton export sales. With rising consumer demand for lion body parts, notably skulls, the export quota presents a further challenge to the industry, regulators and conservationists alike, with 52% of respondents indicating they would adapt by seeking 'alternative markets' for lion bones if the export quota allocation restricted their business. Recognizing that trade policy toward large carnivores represents a 'wicked problem', we anticipate that these results will inform future deliberations, which must nonetheless also be informed by challenging inclusive engagements with all relevant stakeholders.

Evaluation of microclimates and assessment of thermal comfort of Panthera leo in the Masai Mara National Reserve, Kenya.

Quantifying comfort levels of lions within the Masai Mara National Reserve in Kenya is the main focus of this study. Its discourse delineates step by step the process of quantifying comfort levels of lions within the Mara. Resource-efficient measures for humans in the built environment have long been developed through the creation of passive zones and modulated ventilation. In an analogous manner, new procedures are being adapted for creating optimized microclimates in natural game reserves. This involves CFD (computational fluid dynamics)-inspired landscaping. It is seen that the predicted mean vote (PMV) values-measures of thermal comfort-exceed the expected comfortable ranges suitable for normal functioning of lions in the reserve. This calls for a detailed exploration on sustainable development of this sanctuary. The paper illustrates how modern tools in computational fluid dynamics can be used along with standard ecological models to ascertain the optimal extent of airflow, levels of hydration, and land use pattern changes affecting the prevailing microclimate.

A genome-wide data assessment of the African lion (Panthera leo) population genetic structure and diversity in Tanzania.

The African lion (Panthera leo), listed as a vulnerable species on the IUCN Red List of Threatened Species (Appendix II of CITES), is mainly impacted by indiscriminate killing and prey base depletion. Additionally, habitat loss by land degradation and conversion has led to the isolation of some subpopulations, potentially decreasing gene flow and increasing inbreeding depression risks. Genetic drift resulting from weakened connectivity between strongholds can affect the genetic health of the species. In the present study, we investigated the evolutionary history of the species at different spatiotemporal scales. Therefore, the mitochondrial cytochrome b gene (N = 128), 11 microsatellites (N = 103) and 9,103 SNPs (N = 66) were investigated in the present study, including a large sampling from Tanzania, which hosts the largest lion population among all African lion range countries. Our results add support that the species is structured into two lineages at the continental scale (West-Central vs East-Southern), underlining the importance of reviewing the taxonomic status of the African lion. Moreover, SNPs led to the identification of three lion clusters in Tanzania, whose geographical distributions are in the northern, southern and western regions. Furthermore, Tanzanian lion populations were shown to display good levels of genetic diversity with limited signs of inbreeding. However, their population sizes seem to have gradually decreased in recent decades. The highlighted Tanzanian African lion population genetic differentiation appears to have resulted from the combined effects of anthropogenic pressure and environmental/climatic factors, as further discussed.

Linking social and spatial networks to viral community phylogenetics reveals subtype-specific transmission dynamics in African lions.

Heterogeneity within pathogen species can have important consequences for how pathogens transmit across landscapes; however, discerning different transmission routes is challenging. Here, we apply both phylodynamic and phylogenetic community ecology techniques to examine the consequences of pathogen heterogeneity on transmission by assessing subtype-specific transmission pathways in a social carnivore. We use comprehensive social and spatial network data to examine transmission pathways for three subtypes of feline immunodeficiency virus (FIVPle ) in African lions (Panthera leo) at multiple scales in the Serengeti National Park, Tanzania. We used FIVPle molecular data to examine the role of social organization and lion density in shaping transmission pathways and tested to what extent vertical (i.e., father- and/or mother-offspring relationships) or horizontal (between unrelated individuals) transmission underpinned these patterns for each subtype. Using the same data, we constructed subtype-specific FIVPle co-occurrence networks and assessed what combination of social networks, spatial networks or co-infection best structured the FIVPle network. While social organization (i.e., pride) was an important component of FIVPle transmission pathways at all scales, we find that FIVPle subtypes exhibited different transmission pathways at within- and between-pride scales. A combination of social and spatial networks, coupled with consideration of subtype co-infection, was likely to be important for FIVPle transmission for the two major subtypes, but the relative contribution of each factor was strongly subtype-specific. Our study provides evidence that pathogen heterogeneity is important in understanding pathogen transmission, which could have consequences for how endemic pathogens are managed. Furthermore, we demonstrate that community phylogenetic ecology coupled with phylodynamic techniques can reveal insights into the differential evolutionary pressures acting on virus subtypes, which can manifest into landscape-level effects.

Numerical assessment in the wild: insights from social carnivores.

Playback experiments have proved to be a useful tool to investigate the extent to which wild animals understand numerical concepts and the factors that play into their decisions to respond to different numbers of vocalizing conspecifics. In particular, playback experiments have broadened our understanding of the cognitive abilities of historically understudied species that are challenging to test in the traditional laboratory, such as members of the Order Carnivora. Additionally, playback experiments allow us to assess the importance of numerical information versus other ecologically important variables when animals are making adaptive decisions in their natural habitats. Here, we begin by reviewing what we know about quantity discrimination in carnivores from studies conducted in captivity. We then review a series of playback experiments conducted with wild social carnivores, including African lions, spotted hyenas and wolves, which demonstrate that these animals can assess the number of conspecifics calling and respond based on numerical advantage. We discuss how the wild studies complement those conducted in captivity and allow us to gain insights into why wild animals may not always respond based solely on differences in quantity. We then consider the key roles that individual discrimination and cross-modal recognition play in the ability of animals to assess the number of conspecifics vocalizing nearby. Finally, we explore new directions for future research in this area, highlighting in particular the need for further work on the cognitive basis of numerical assessment skills and experimental paradigms that can be effective in both captive and wild settings.This article is part of a discussion meeting issue 'The origins of numerical abilities'.

Sustainability and Long Term-Tenure: Lion Trophy Hunting in Tanzania.

It is argued that trophy hunting of large, charismatic mammal species can have considerable conservation benefits but only if undertaken sustainably. Social-ecological theory suggests such sustainability only results from developing governance systems that balance financial and biological requirements. Here we use lion (Panthera leo) trophy hunting data from Tanzania to investigate how resource ownership patterns influence hunting revenue and offtake levels. Tanzania contains up to half of the global population of free-ranging lions and is also the main location for lion trophy hunting in Africa. However, there are concerns that current hunting levels are unsustainable. The lion hunting industry in Tanzania is run by the private sector, although the government leases each hunting block to companies, enforces hunting regulation, and allocates them a species-specific annual quota per block. The length of these leases varies and theories surrounding property rights and tenure suggest hunting levels would be less sustainable in blocks experiencing a high turnover of short-term leases. We explored this issue using lion data collected from 1996 to 2008 in the Selous Game Reserve (SGR), the most important trophy hunting destination in Tanzania. We found that blocks in SGR with the highest lion hunting offtake were also those that experienced the steepest declines in trophy offtake. In addition, we found this high hunting offtake and the resultant offtake decline tended to be in blocks under short-term tenure. In contrast, lion hunting levels in blocks under long-term tenure matched more closely the recommended sustainable offtake of 0.92 lions per 1000 km2. However, annual financial returns were higher from blocks under short-term tenure, providing $133 per km2 of government revenue as compared to $62 per km2 from long-term tenure blocks. Our results provide evidence for the importance of property rights in conservation, and support calls for an overhaul of the system in Tanzania by developing competitive market-based approaches for block allocation based on long-term tenure of ten years.

Conservation and monitoring of a persecuted African lion population by Maasai warriors.

Although Africa has many threatened species and biological hot spots, there are few citizen science schemes, particularly in rural communities, and there has been limited evaluation of existing programs. We engaged traditional Maasai warriors (pastoralist men aged 15 to 35) in community-based conservation and demographic monitoring of a persecuted African lion (Panthera leo) population. Through direct engagement, we investigated whether a citizen science approach employing local warriors, who had no formal education, could produce reliable data on the demographics, predation, and movements of a species with which their communities have been in conflict for generations. Warriors were given benefits such as literacy training and skill enhancement and engaged in the monitoring of the lions. The trained warriors reported on lion sign across an area nearly 4000 km(2) . Scientists worked together with the warriors to verify their reports and gather observations on the lion population. Using the verified reports and collected observations, we examined our scientific knowledge relative to the lion population preceding and during the citizen science program. Our observations showed that data quality and quantity improved with the involvement and training of the participants. Furthermore, because they engaged in conservation and gained personal benefits, the participants came to appreciate a species that was traditionally their foe. We believe engaging other local communities in biodiversity conservation and monitoring may be an effective conservation approach in rural Africa.

Understanding tourists' preference for mammal species in private protected areas: is there a case for extralimital species for ecotourism?

Private Protected Areas (PPAs) often use wildlife-based ecotourism as their primary means of generating business. Achieving tourist satisfaction has become a strong driving goal in the management of many PPAs, often at the expense of biodiversity. Many extralimitral species, those which historically did not occur in an area, are stocked in PPAs with the intention of increasing ecotourism attractions. Even though the ecological and economic costs of stocking these species are high, the social benefits are not understood and little information exists globally on the ecotourism role of extralimital species. This study assessed the value of stocking extralimital species using questionnaire-based surveys and observing tourists in Shamwari Private Game Reserve in the Eastern Cape Province of South Africa. No difference was found between indigenous and extralimital species with regards to the tourists' weighted scoring system, average amount tourists were willing to pay, total viewing time, average viewing time or the likelihood of stopping to view species when encountered on game drives. During game drives a strong preference was found for the elephant (Loxodonta africana), lion (Panthera leo), leopard (Panthera pardus) and cheetah (Acynonix jubatus). With the exception of the cheetah, these species are all members of the "big five" and are indigenous. Species availability and visibility, however, may influence the amount of time tourists spend at an animal sighting. Our analysis suggests that certain extralimital species (typically larger and charismatic species) contribute to tourist satisfaction, while particularly the smaller extralimital species add little to the game viewing experience, but add to the costs and risks of the PPAs. We recommend that extralimital species introductions for ecotourism purposes should be approached with caution with regards to the risks to the sustainability of PPAs.

Efficacy of two lion conservation programs in Maasailand, Kenya.

Lion (Panthera leo) populations are in decline throughout most of Africa. The problem is particularly acute in southern Kenya, where Maasai pastoralists have been spearing and poisoning lions at a rate that will ensure near term local extinction. We investigated 2 approaches for improving local tolerance of lions: compensation payments for livestock lost to predators and Lion Guardians, which draws on local cultural values and knowledge to mitigate livestock-carnivore conflict and monitor carnivores. To gauge the overall influence of conservation intervention, we combined both programs into a single conservation treatment variable. Using 8 years of lion killing data, we applied Manski's partial identification approach with bounded assumptions to investigate the effect of conservation treatment on lion killing in 4 contiguous areas. In 3 of the areas, conservation treatment was positively associated with a reduction in lion killing. We then applied a generalized linear model to assess the relative efficacy of the 2 interventions. The model estimated that compensation resulted in an 87-91% drop in the number of lions killed, whereas Lion Guardians (operating in combination with compensation and alone) resulted in a 99% drop in lion killing.

The lion in West Africa is critically endangered.

The African lion has declined to <35,000 individuals occupying 25% of its historic range. The situation is most critical for the geographically isolated populations in West Africa, where the species is considered regionally endangered. Elevating their conservation significance, recent molecular studies establish the genetic distinctiveness of West and Central African lions from other extant African populations. Interventions to save West African lions are urgently required. However formulating effective conservation strategies has been hampered by a lack of data on the species' current distribution, status, and potential management deficiencies of protected areas (PAs) harboring lions. Our study synthesized available expert opinion and field data to close this knowledge gap, and formulate recommendations for the conservation of West African lions. We undertook lion surveys in 13 large (>500 km²) PAs and compiled evidence of lion presence/absence for a further eight PAs. All PAs were situated within Lion Conservation Units, geographical units designated as priority lion areas by wildlife experts at a regional lion conservation workshop in 2005. Lions were confirmed in only 4 PAs, and our results suggest that only 406 (273-605) lions remain in West Africa, representing <250 mature individuals. Confirmed lion range is estimated at 49,000 km², or 1.1% of historical range in West Africa. PAs retaining lions were larger than PAs without lions and had significantly higher management budgets. We encourage revision of lion taxonomy, to recognize the genetic distinctiveness of West African lions and highlight their potentially unique conservation value. Further, we call for listing of the lion as critically endangered in West Africa, under criterion C2a(ii) for populations with <250 mature individuals. Finally, considering the relative poverty of lion range states in West Africa, we call for urgent mobilization of investment from the international community to assist range states to increase management effectiveness of PAs retaining lions.

Insights into the management of large carnivores for profitable wildlife-based land uses in African savannas.

Large African predators, especially lions (Panthera leo) and leopards (Panthera pardus), are financially valuable for ecotourism and trophy hunting operations on land also utilized for the production of other wildlife species for the same purpose. Predation of ungulates used for trophy hunting can create conflict with landholders and trade off thus exists between the value of lions and leopards and their impact on ungulate populations. Therefore productionist and conservation trade-offs are complexly graded and difficult to resolve. We investigated this with a risk-benefit analysis on a large private wildlife production area in Zimbabwe. Our model showed that lions result in substantial financial costs through predation on wild ungulates that may not be offset by profits from hunting them, whereas the returns from trophy hunting of leopards are projected to exceed the costs due to leopard predation. In the absence of additional income derived from photo-tourism the number of lions may need to be managed to minimize their impact. Lions drive important ecological processes, but there is a need to balance ecological and financial imperatives on wildlife ranches, community wildlife lands and other categories of multiple use land used for wildlife production. This will ensure the competitiveness of wildlife based land uses relative to alternatives. Our findings may thus be limited to conservancies, community land-use areas and commercial game ranches, which are expansive in Africa, and should not necessarily applied to areas where biodiversity conservation is the primary objective, even if hunting is allowed there.

Conserving large carnivores: dollars and fence.

Conservationists often advocate for landscape approaches to wildlife management while others argue for physical separation between protected species and human communities, but direct empirical comparisons of these alternatives are scarce. We relate African lion population densities and population trends to contrasting management practices across 42 sites in 11 countries. Lion populations in fenced reserves are significantly closer to their estimated carrying capacities than unfenced populations. Whereas fenced reserves can maintain lions at 80% of their potential densities on annual management budgets of $500 km(-2) , unfenced populations require budgets in excess of $2000 km(-2) to attain half their potential densities. Lions in fenced reserves are primarily limited by density dependence, but lions in unfenced reserves are highly sensitive to human population densities in surrounding communities, and unfenced populations are frequently subjected to density-independent factors. Nearly half the unfenced lion populations may decline to near extinction over the next 20-40 years.

A molecular analysis of African lion (Panthera leo) mating structure and extra-group paternity in Etosha National Park.

The recent incorporation of molecular methods into analyses of social and mating systems has provided evidence that mating patterns often differ from those predicted by group social organization. Based on field studies and paternity analyses at a limited number of sites, African lions are predicted to exhibit a strict within-pride mating system. Extra-group paternity has not been previously reported in African lions; however, observations of extra-group associations among lions inhabiting Etosha National Park in Namibia suggest deviation from the predicted within-pride mating pattern. We analysed variation in 14 microsatellite loci in a population of 164 African lions in Etosha National Park. Genetic analysis was coupled with demographic and observational data to examine pride structure, relatedness and extra-group paternity (EGP). EGP was found to occur in 57% of prides where paternity was analysed (n = 7), and the overall rate of EGP in this population was 41% (n = 34). Group sex ratio had a significant effect on the occurrence of EGP (P < 0.05), indicating that variation in pride-level social structure may explain intergroup variation in EGP. Prides with a lower male-to-female ratio were significantly more likely to experience EGP in this population. The results of this study challenge the current models of African lion mating systems and provide evidence that social structure may not reflect breeding structure in some social mammals.