Lions,trophy hunting and beyond: knowledge gaps and why they matter

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.     

In the absence of a “landscape of fear”: How lions,hyenas, and cheetahs coexist

Aggression by top predators can create a “landscape of fear” in which subordinate predators restrict their activity to low‐risk areas or times of day. At large spatial or temporal scales, this can result in the costly loss of access to resources. However, fine‐scale reactive avoidance may minimize the risk of aggressive encounters for subordinate predators while maintaining access to resources, thereby providing a mechanism for coexistence. We investigated fine‐scale spatiotemporal avoidance in a guild of African predators characterized by intense interference competition. Vulnerable to food stealing and direct killing, cheetahs are expected to avoid both larger predators; hyenas are expected to avoid lions. We deployed a grid of 225 camera traps across 1,125 km² in Serengeti National Park, Tanzania, to evaluate concurrent patterns of habitat use by lions, hyenas, cheetahs, and their primary prey. We used hurdle models to evaluate whether smaller species avoided areas preferred by larger species, and we used time‐to‐event models to evaluate fine‐scale temporal avoidance in the hours immediately surrounding top predator activity. We found no evidence of long‐term displacement of subordinate species, even at fine spatial scales. Instead, hyenas and cheetahs were positively associated with lions except in areas with exceptionally high lion use. Hyenas and lions appeared to actively track each, while cheetahs appear to maintain long‐term access to sites with high lion use by actively avoiding those areas just in the hours immediately following lion activity. Our results suggest that cheetahs are able to use patches of preferred habitat by avoiding lions on a moment‐to‐moment basis. Such fine‐scale temporal avoidance is likely to be less costly than long‐term avoidance of preferred areas: This may help explain why cheetahs are able to coexist with lions despite high rates of lion‐inflicted mortality, and highlights reactive avoidance as a general mechanism for predator coexistence.     

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.     

Socio‐demographic factors influence the attitudes of local residents towards trophy hunting activities in the Kilombero Valley,Tanzania

Within the Kilombero Game Controlled Area (KGCA) of Tanzania, protection is offered to large mammal populations by trophy hunting concessions that maintain natural habitat through the prevention of extensive human encroachment. The opinions of local communities to wildlife management operations such as trophy hunting play an important role in their long‐term viability. This study addresses the influence of socio‐demographic factors on the opinions of local communities to trophy hunting in areas that are not part of community‐based management projects, which is where most research of this type has previously been conducted. Semi‐structured questionnaires were conducted in 24 villages within the Kilombero Valley (fifteen interviews per village) in August–December 2007. The extent to which socio‐demographic factors including location (e.g. village of residence) and individual respondent characteristics (e.g. gender and age) influenced the respondents’ opinions was analysed. Of these socio‐demographic factors, all, except age and district of residence, were found to influence the opinions of local residents. Socio‐demographic factors play an important role in determining local communities’ attitudes towards trophy hunting, and this must be taken into account during the design and assessment of wildlife management conservation strategies, both locally in the KGCA and in similar national and international initiatives.     

Using Mountain Lion Habitat Selection in Management

Wildlife agencies are generally tasked with managing and conserving species at state and local levels simultaneously. Thus, it is necessary for wildlife agencies to understand basic ecological processes of a given species at multiple scales to aid decision making at commensurately varied spatial and behavioral scales. Mountain lions (Puma concolor) occur throughout California, USA, and are at the center of a variety of management and conservation issues. For example, they are genetically and demographically at risk in 1 region yet apparently stable and negatively affecting endangered species in another. Currently, no formal plan exists for mountain lions in California to deal with these diverse scenarios involving issues of local mountain lion population viability and problems related to predation of endangered species. To facilitate development of a state-wide management and conservation plan, we quantified habitat selection by mountain lions at 2 spatial scales across the range of environmental conditions in which the species is found in California. Our analyses used location data from individuals (n = 263) collared across the state from 2001–2019. At the home range scale, mountain lions selected habitat to prioritize meeting energetic demands. At the within home range scale, mountain lions avoided areas of human activity. Further, our analyses revealed 165,350–170,085 km², depending on season, of suitable mountain lion habitat in California. Fifty percent of the suitable habitat was on unprotected lands and thus vulnerable to development. These habitat selection models will help in the development of a state-wide conservation and management plan for mountain lions in California by guiding mountain lion population monitoring through time, prioritization of habitat to be conserved for maintaining demographic connectivity and gene flow, and efforts to mediate mountain lion-prey interactions. Our work and application area will help with wildlife policy and management decisions related to depredation problems at the local scale and issues of habitat connectivity at the statewide scale. © 2019 The Wildlife Society.     

A man-eating lion (Panthera leo) from Tanzania with a toothache

Between August 2002 and April 2004, a man-eating lion killed 35 people and injured at least 9 in a 350-km² area 150 km south-west of Dar es Salaam in Tanzania. Man-eating by lions is common in Southern Tanzania, but this case was exceptional due to the high number of victims attributed to a single animal. After having been killed by game scouts and villagers, it was found to be a young adult male. Dental examination revealed a broken upper left molar where a serious abscess and caries had developed. The lion must have been in permanent pain, and this probably explains its preference for man-eating. Most man-eating lions in Tanzania are healthy animals without signs of infirmity.     

The impact of tourist hunting on large mammals in Tanzania: an initial assessment

In Tanzania, where tourist hunting is employed as a conservation tool for habitat protection, information on population sizes and hunting offtake was used to assess the impact of tourist hunting on mammal densities. In general, tourist hunting pressure was unrelated to local population sizes, but for most species, animals were removed at a level of less than 10% of the local population size, suggesting that over-exploitation was unlikely. Eland, however, and perhaps small antelope, bushbuck, kudu and reedbuck were hunted at levels which may be unsustainable in the long term. Analyses also identified areas of Tanzania with high levels of tourist hunting pressure, showed that, in certain areas, species with small population sizes such as eland could be declining as a result of tourist hunting, and suggested that current levels of lion and leopard offtake are too high. These findings, although preliminary, allow recommendations to be put forward for changing hunting quotas for certain species in particular areas of Tanzania.     

Hunting success of lions affected by the moon's phase in a wooded habitat

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.     

Estimating past and future male loss in three Zambian lion populations

African lions (Panthera leo) are declining continent-wide, with protected area populations subject to a variety of anthropogenic effects. Zambia contains viable lion populations of considerable importance for photographic and hunting tourism, but long-term lion demographic data do not exist to guide recent management directives and population projections under different strategies. We described population size, as well as age and sex structure of lions in 3 Zambian national park populations bordering hunting areas, and found them to be male-depleted relative to other systems. We then estimated rates of adult male loss leading to male depletion in these populations and the effect of different future hunting management options on population characteristics. Predictions from matrix population models constructed within a Bayesian framework confirmed that the observed population structure was likely due to high rates of adult male loss and that instituting age limits on male harvests with quota reductions would reduce male depletion, improve tourism by providing older and more abundant males, and slightly increase population size. Reducing male mortality from wire snare poaching would also result in similar demographic impacts, and in concert with changes in hunting regulations would substantially improve the quality and quantity of adult male lions. However, model results varied depending on whether we assumed historical population stability. Predictions assuming negative historical growth rate indicated that substantially more conservative lion harvest management is warranted. We discuss the relevance of these findings for maintaining viable lion populations in and around protected areas in Zambia. © 2012 The Wildlife Society.     

Fear of darkness, the full moon and the nocturnal ecology of African lions

Nocturnal carnivores are widely believed to have played an important role in human evolution, driving the need for night-time shelter, the control of fire and our innate fear of darkness. However, no empirical data are available on the effects of darkness on the risks of predation in humans. We performed an extensive analysis of predatory behavior across the lunar cycle on the largest dataset of lion attacks ever assembled and found that African lions are as sensitive to moonlight when hunting humans as when hunting herbivores and that lions are most dangerous to humans when the moon is faint or below the horizon. At night, people are most active between dusk and 10:00 pm, thus most lion attacks occur in the first weeks following the full moon (when the moon rises at least an hour after sunset). Consequently, the full moon is a reliable indicator of impending danger, perhaps helping to explain why the full moon has been the subject of so many myths and misconceptions.     

Methods for lion monitoring: a comparison from the Selous Game Reserve,Tanzania

The Selous Game Reserve in Tanzania is believed to contain Africa's largest population of lions (Panthera leo), making it a popular destination for trophy hunters and photographic tourists. However, a lack of recent data has raised concerns about the conservation status of this iconic population, so we collected two types of population data between 2006 and 2009. First, we identified 112 individual animals in an 800 km² study area in the photographic tourism part of Selous, giving a density of 0.14 individuals km^?2. This density estimate was similar to results using the same method from 1997 to 1999, but the adult sex ratio has decreased from 1 male : 1.3 female in 1997 to 1 male : 3 females in 2009. Second, using buffalo calf distress calls, we conducted call‐up surveys to census lions in three hunting sectors in the west, east and south of Selous and in the northern photographic area. Estimated adult lion densities varied from 0.02 to 0.10 km^?2, allowing an overall population estimate of 4,300 (range: 1,700–6,900). Our results highlight the value of call‐ups in surveying cryptic hunted carnivores but stress the importance of long‐term projects for calibrating the responses to call‐ups and for measuring trends in demography and population size.     

Genetic structure of mountain lion (Puma concolor) populations in California

Analysis of 12 microsatellite loci from431 mountain lions (Puma concolor)revealed distinct genetic subdivision that wasassociated with geographic barriers andisolation by distance in California. Levels ofgenetic variation differed among geographicregions, and mountain lions that inhabitedcoastal areas exhibited less heterozygositythan those sampled inland. The San FranciscoBay and Sacramento-San Joaquin River Delta, theCentral Valley, and the Los Angeles Basinappeared to be substantial barriers to geneflow, and allele frequencies of populationsseparated by those features differedsubstantially. A partial barrier to gene flowappeared to exist along the crest of the SierraNevada. Estimated gene flow was high amongmountain lions inhabiting the Modoc Plateau,the western Sierra Nevada, and northern sectionof the eastern Sierra Nevada. SouthernCalifornia mountain lion populations mayfunction as a metapopulation; however, humandevelopments threaten to eliminate habitat andmovement corridors. While north-south geneflow along the western Sierra Nevada wasestimated to be very high, projected loss andfragmentation of foothill habitat may reducegene flow and subdivide populations. Preservation of existing movement corridorsamong regions could prevent population declinesand loss of genetic variation. This studyshows that mountain lion management andconservation efforts should be individualizedaccording to region and incorporatelandscape-level considerations to protecthabitat connectivity.     

南非Hluhluwe-Umfolozi公园植被对非洲狮群大小的影响

非洲狮(Pantheraleo)群处于不断的分裂-聚合过程,其亚群大小与原狮群大小以及诸如狮群捕食成功度、协作幼仔保护及狮群领域防卫等社会性因素有关。我们分析了南非一个保护区(Hluhluwe-Umfolozi公园)15年间的狮群大小,其成年雌狮和成年雄狮群的大小在保护区内呈显著性变化。在开放生境中雌狮结大群,而茂密植被生境中则结小群。雄狮群大小与植被类型无关,但在大的雌狮群分布区域发现大雄狮群。令人惊奇的是,作为幼狮成活率指标的幼狮群大小和雌狮平均幼仔数在保护区内没有显著性差异。我们认为,雌狮群大小的不同是对植被结构的适应,即在开放生境中大群有利于协作狩猎、育幼、共同保护幼仔、应对高强度群间领域防卫竞争。相反,茂密植被中,群间竞争很低,高密度植被覆盖有利于狩猎成功及补偿小狮群对幼仔保护的不足。雌狮平均幼仔数在整个保护区内没有变化,我们认为是由于开放生境和茂密植被对狮群大小起到相似的调节作用     

QUANTIFICATION OF TERRESTRIAL HAUL-OUT AND ROOKERY CHARACTERISTICS OF STELLER SEA LIONS

Steller sea lions ( Eumetopias jubatus ) are known to have occupied the same terrestrial haul-out and rookery sites across the North Pacific Rim for centuries, but it is not known why they choose and stay at these locations, or what defines their preferred habitat. Classifying and comparing the shoreline type of haul-outs and rookeries against sites not used by Steller sea lions showed that they preferentially locate their haul-outs and rookeries on exposed rocky shorelines and wave-cut platforms. However, no preference was found for selecting rookeries on sheltered shore types. Shoreline types used less frequently by sea lions included fine-to-medium-grained sand beaches, mixed sand and gravel beaches, gravel beaches, and sheltered rocky shores. Quantifying the shoreline types used by sea lions confirms anecdotal reports of habitat preferences and may prove useful in identifying and protecting sea lion terrestrial habitat, or in forecasting how climate change might affect the distribution of sea lions.     

Depth and movement behaviour of the Pacific sleeper shark in the north-east Pacific Ocean

Satellite-linked radio telemetry was used to study the geographic movements and vertical movement behaviour of the Pacific sleeper shark Somniosus pacificus . The fish were tagged near Steller sea lion Eumetopias jubatus rookeries in the Gulf of Alaska during periods when Steller sea lions pups were most vulnerable to predation; when Steller sea lion pups first enter the water (July to August) and when Steller sea lion pups are weaned (April to May). Final locations recovered from most Pacific sleeper sharks (76%) were within 100 km of release locations, 16% were within 100–250 km and 8% were within 250–500 km. The most striking behavioural feature was their extensive, nearly continuous vertical movements. Median daily depth range was 184 m; the most time (61%) was spent between 150 and 450 m, but ascents above 100 m were common (58% of days). Median vertical movement rate was 6 km day⁻¹ and steady. The longest period of continuous vertical movement (> 60 m h⁻¹) was 330 h. Systematic vertical oscillations were most common (60%), followed by diel vertical migrations (25%) and irregular vertical movements (15%). The Pacific sleeper sharks travelled below the photic zone during the day and approached the surface at night. Pacific sleeper sharks appear to employ a stealth and ambush hunting strategy that incorporates slow vertical oscillations to search for prey, and cryptic colouration and cover of darkness to avoid detection by potential prey. The depth and geographic range of Pacific sleeper shark and Steller sea lions overlap near four important Steller sea lion rookeries in the northern Gulf of Alaska, so the potential exists for predation to occur. None of the tissues in the stomachs of the 198 Pacific sleeper sharks collected during a companion diet study, however, were identified as Steller sea lion.     

Assessing trophy hunting in South Africa by comparing hunting and exporting databases

Trophy hunting constitutes a major part of the global wildlife tourism industry and is connected through the export of kills to international wildlife trade. Inconsistencies between kills and exports can contribute to identifying illegal trophy hunting that constitutes a major threat to biodiversity conservation. This paper quantitatively analyses to what extent the data of trophy hunting kills and of trophy exports are consistent using the example of South Africa. Data was extracted from two different sources for the year 2018. These sources were trade data reported under the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES) and its Trade Database on the export of trophy items from South Africa, which is compared with the South African Professional Hunting statistics (SAPHs) containing trophy kills information for each species and the country of origin of the hunter. The data of trophy hunting kills and data of trophy hunting export was found to differ to varying degrees across the countries contributing to trophy hunting and CITES-listed trophy-hunted species. We found that both databases report hunting of the same 28 threatened taxa. On the other hand, the same data reports that hunters of 64 nationalities participated in a total of 4,726 trophy kills, while only 3,131 hunting trophy items were exported from South Africa, to 37 countries as final destinations. Among the possible reasons for the discrepancies found, we suggest that the time required to taxidermize trophy specimens may delay the items being addressed to their final destination, in addition to the dual citizenship of some hunters that should also have some influence on our results, as well as the important commercial destination of Brussels Airport. The USA show the highest absolute number of trophy hunters, followed by Spain, but Denmark is the highest driver per capita. Therefore, greater participation of these countries in conservation policies for endangered species is necessary. Additionally, a more detailed differentiation of the term “trophy” to more specific terms such as claws, skins, skulls, etc. may improve reporting systems to easier identify illegal activities related to hunting.     

‘Site’ing the right reasons: critical evaluation of conservation planning for the Asiatic lion

The resurrection of Asiatic lions (Panthera leo persica) from the brink of extinction is a remarkable conservation success story. Yet, occurrence of lions as a single population makes them vulnerable to extinction from genetic and environmental factors. Asiatic lions exist as a single free ranging population of 360 individuals in Gir Protected Area (PA; about 290 lions) and surrounding satellite areas (68 lions), namely Girnar Wildlife Sanctuary (WLS), coastal areas, hill ranges extending from Mitiyala-Savarkundla-Palitana-Shihor spreading across Junagadh, Amreli and Bhavnagar districts of Gujarat State, India. This paper traces the conservation history, current conservation pressures, and critically evaluates current conservation planning based on lion ecology and existing anthropogenic pressures. Conservation proposals for translocation of lions to alternate habitats in Kuno WLS and Barda WLS are awaiting final implementation. An alternate initiative is aimed at developing satellite lion habitats, improving corridor connectivity, and facilitating natural dispersal and expansion of lion habitats. The paper evaluates human population living within 2 km boundary of lion habitats and between Gir PA and satellite habitats to show that dispersing lions have to cross heavily populated habitations taking refuge in agriculture fields and scattered forest patches. Satellite habitats vary in size from 18 km² (Mitiyala WLS) to 250 km² (scattered forests Hipavadli-Savarkundla-Palitana zone along Shetrunji river) and are inadequate to maintain natural ranging and movement requirements of territorial lions. These habitats are varied in vegetation, terrain, human pressure, and distance to source population. Though satellite lion habitats are important suboptimal habitats for dispersing lions, long-term conservation planning require planned restoration of mosaic habitats for growing populations.     

Can an herbivore affect where a top predator kills its prey by modifying woody vegetation structure?

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, < 150 m, n = 167). Elephant-impacted sites had a slightly higher visibility and a longer distance to a potential ambush site than non-impacted sites, but these relationships were characterized by a high variability. At large scale, kill sites were characterized by higher levels of elephant impacts compared to random sites. At microhabitat scale, compared to the direct nearby available habitat, kill sites were characterized by a reduced distance to a potential ambush site. We suggest a conceptual framework whereby the relative importance of habitat features and prey abundance could change upon the scale considered.     

Steller sea lion spatial‐use patterns derived from a Bayesian model of opportunistic observations

Despite acquisition of a substantial catalog of telemetry data from Steller sea lions (Eumetopias jubatus) over the past two decades, scientists still lack comprehensive regionally explicit knowledge about Steller sea lion habitat use. The Platforms of Opportunity data contain records of Steller sea lion sightings throughout the species’ entire range and have potential to fill gaps in knowledge about their spatial use; however, the data have not previously been used because effort (e.g., time spent surveying or area sampled) was not recorded when sightings were obtained. For this study a novel approach was used to overcome the lack of effort data through development of an effort index and a Bayesian negative binomial model. The model quantified Steller sea lion encounter rates and associated uncertainty within 15 × 15 km² grid cells across the species’ entire range. Year‐round, as well as breeding and nonbreeding season encounter rates were estimated. The results of this analysis identify several previously undocumented areas of high use by Steller sea lions, indicate that only 37% of Steller sea lion high‐use areas fall within designated critical habitat, and demonstrate that use of depth and distance from shore as indicators of Steller sea lion habitat is contraindicated.     

Trophy hunting and perceived risk in closed ecosystems: Flight behaviour of three gregarious African ungulates in a semi‐arid tropical savanna

Although being an important conservation tool in Africa, trophy hunting is known to influence risk perception in wildlife species, thus affecting the behaviour and fitness of most targeted species. We studied the effects of trophy hunting on the flight behaviour of impala (Aepyceros melampus), greater kudu (Tragelaphus strepsiceros) and sable (Hippotragus niger) in two closed ecosystems, Cawston Ranch (hunting area) and Stanley and Livingstone Private Game Reserve (tourist area), western Zimbabwe. Using standardized field procedures, we assessed the flight behavioural responses of the three species in two seasons: non‐hunting (December–March) and hunting (April–November) between March 2013 and November 2014. We tested the effect of habitat, group size, sex, season, start distance and alert distance on flight initiation distance using linear mixed models. Habitat, group size sex and alert distance did not have any effect on flight initiation distance for the three species. The three species were more alert and displayed longer flight initiation distances in the hunting area compared with the tourist area. Flight initiation distances for the three species were higher during the hunting season for the hunting area and low during the non‐hunting season. Flight distances of the three species did not differ between the hunting area and the tourist area. We concluded that trophy hunting increased perceived risk of wild ungulates in closed hunting areas, whereas ungulates in non‐hunting areas are less responsive and somehow habituated to human presence. Management plans should include minimum approach distances by tourists as well as establishing seasonal restrictions on special zones to promote species viability. Research aimed at integrating behavioural responses with physiological aspects of target species should be promoted to ensure that managers are able to deal with the behavioural trade‐offs of trophy hunting at local and regional scale.