The significance of African lions for the financial viability of trophy hunting and the maintenance of wild land

Recent studies indicate that trophy hunting is impacting negatively on some lion populations, notably in Tanzania. In 2004 there was a proposal to list lions on CITES Appendix I and in 2011 animal-welfare groups petitioned the United States government to list lions as endangered under their Endangered Species Act. Such listings would likely curtail the trophy hunting of lions by limiting the import of lion trophies. Concurrent efforts are underway to encourage the European Union to ban lion trophy imports. We assessed the significance of lions to the financial viability of trophy hunting across five countries to help determine the financial impact and advisability of the proposed trade restrictions. Lion hunts attract the highest mean prices (US$24,000-US$71,000) of all trophy species. Lions generate 5-17% of gross trophy hunting income on national levels, the proportional significance highest in Mozambique, Tanzania, and Zambia. If lion hunting was effectively precluded, trophy hunting could potentially become financially unviable across at least 59,538 km(2) that could result in a concomitant loss of habitat. However, the loss of lion hunting could have other potentially broader negative impacts including reduction of competitiveness of wildlife-based land uses relative to ecologically unfavourable alternatives. Restrictions on lion hunting may also reduce tolerance for the species among communities where local people benefit from trophy hunting, and may reduce funds available for anti-poaching. If lion off-takes were reduced to recommended maximums (0.5/1000 km(2)), the loss of viability and reduction in profitability would be much lower than if lion hunting was stopped altogether (7,005 km(2)). We recommend that interventions focus on reducing off-takes to sustainable levels, implementing age-based regulations and improving governance of trophy hunting. Such measures could ensure sustainability, while retaining incentives for the conservation of lions and their habitat from hunting.     

Does captivity influence territorial and hunting behaviour? Assessment for an ex situ reintroduction program of African lions Panthera leo

Maintaining a territory and being able to hunt are imperative for the success of African lion Panthera leo prides. We aimed to determine whether captive‐origin prides display similar territorial and hunting behaviour to wild lions. Behaviours and locations of two captive‐origin prides and one wild pride were collected through direct observation. All prides established territories, and core areas corresponded to resource requirements. There was no evidence that pride origin affected territorial or hunting behaviour. Captive‐origin prides exhibited behaviours that lead us to be optimistic about each pride's ability to establish and defend a territory successfully, and to hunt, following reintroduction.     

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.     

‘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.     

The Effect of Moonlight on Scopoli's Shearwater Calonectris diomedea Colony Attendance Patterns and Nocturnal Foraging: A Test of the Foraging Efficiency Hypothesis

Moonlight is known to affect the nocturnal behaviour and activity rhythms of many organisms. For instance, predators active at night may take advantage from increased visibility afforded by the moon, while prey might regulate their activity patterns to become less detectable. Many species of pelagic seabirds attend their colony only at night, in complete darkness, avoiding approaching their nest sites under moonlight. This behaviour has been most often interpreted as an antipredator adaptation (‘predation avoidance’ hypothesis). However, it may also reflect a lower foraging efficiency during moonlit nights (‘foraging efficiency’ hypothesis). Indeed, moonlight may reduce prey availability because preferred seabird prey is known to occur at higher depths in moonlit nights. Using high‐accuracy behavioural information from data loggers, we investigated the effect of moonlight on colony attendance and at‐sea nocturnal foraging in breeding Scopoli's shearwaters Calonectris diomedea. We found that birds departing for self‐feeding trips around the full moon performed longer trips than those departing around the new moon. On nights when the moon was present only partly, nest burrow entrances took place largely in the moonless portion of the night. Moreover, contrary to predictions from the ‘foraging efficiency’ hypothesis, nocturnal foraging activity increased according to moonlight intensity, suggesting that birds increased their foraging activity when prey became more detectable. This study strengthens the idea that colony attendance behaviour is strictly controlled by moonlight in shearwaters, which is possibly related to the perception of a predation risk.     

Seasonal Diet and Prey Preference of the African Lion in a Waterhole-Driven Semi-Arid Savanna

Large carnivores inhabiting ecosystems with heterogeneously distributed environmental resources with strong seasonal variations frequently employ opportunistic foraging strategies, often typified by seasonal switches in diet. In semi-arid ecosystems, herbivore distribution is generally more homogeneous in the wet season, when surface water is abundant, than in the dry season when only permanent sources remain. Here, we investigate the seasonal contribution of the different herbivore species, prey preference and distribution of kills (i.e. feeding locations) of African lions in Hwange National Park, Zimbabwe, a semi-arid African savanna structured by artificial waterholes. We used data from 245 kills and 74 faecal samples. Buffalo consistently emerged as the most frequently utilised prey in all seasons by both male (56%) and female (33%) lions, contributing the most to lion dietary biomass. Jacobs’ index also revealed that buffalo was the most intensively selected species throughout the year. For female lions, kudu and to a lesser extent the group “medium Bovidae” are the most important secondary prey. This study revealed seasonal patterns in secondary prey consumption by female lions partly based on prey ecology with browsers, such as giraffe and kudu, mainly consumed in the early dry season, and grazers, such as zebra and suids, contributing more to female diet in the late dry season. Further, it revealed the opportunistic hunting behaviour of lions for prey as diverse as elephants and mice, with elephants taken mostly as juveniles at the end of the dry season during droughts. Jacobs’ index finally revealed a very strong preference for kills within 2 km from a waterhole for all prey species, except small antelopes, in all seasons. This suggested that surface-water resources form passive traps and contribute to the structuring of lion foraging behaviour.     

A mass capture technique for lions

During a 13 month survey over 600 lions were lured to capture sites and of these 488 were captured and 409 marked in 158 effort nights (two teams worked for 79 nights). Lions were captured in 96 effort nights out of the 158. Groups of up to 16 were captured per effort night with 80% of all observed lions captured. Seasonal capture rates indicated that there was an inverse relationship between rainfall and capture success, the greatest success being recorded during the dry season. Lions were lured to a game carcass with tape recordings of lions and spotted hyaena calls, and scent trails. All lions were darted at night from the back of a parked vehicle. Record ings were audible to 7 km, the effective luring range with regard to lions being about 4 km. Lions and hyaenas also responded to the sounds in the absence of a carcass or scent trails and approached the sound at a brisk walk. Playing roars often caused lions to move away. Recordings used in conjunction with a carcass and scent trails reduced the time lions took to arrive by a factor of 2.7 when compared to using only a carcass and scent trails. Eighty-three per cent of the darted lions remained within 10 m of the carcass. Both sexes and all age classes except cubs under about 1 month were captured by the technique. The mortality rate was 0.82%.     

Seed Dispersal by Golden-headed Lion Tamarins Leontopithecus chrysomelas in Southern Bahian Atlantic Forest, Brazil

Seed dispersal by small primates may be particularly relevant in areas where populations of larger frugivores have been reduced or extinguished by hunting and/or habitat disturbance. In this context, the aim of this study was to evaluate the role of the golden-headed lion tamarin Leontopithecus chrysomelas as a seed disperser in Atlantic forest remnants in Brazil. To this end, we opportunistically collected feces deposited during observations on the feeding behavior of two groups of golden-headed lion tamarins ranging in the degraded areas of the Una Biological Reserve, Bahia, Brazil, from February 2006 to January 2007. We collected 587 fecal samples, of which 524 contained seeds from 24 plant species, distributed over 13 families. Disregarding seeds of<3 mm, the majority of seeds recovered were bromeliad seeds. In general, ingestion of seeds by golden-headed lion tamarins did not improve the germination proportion or decrease the germination delay of seeds, with the exception of Aechmea spp. seeds. The tamarins encountered different habitats during their daily activity period, while feeding and defecating. Consequently, some seeds were transported to different habitats including disturbed areas. Thus, the role of seed dispersal in combination with the daily movement pattern of L. chrysomelas contributes to the persistence of fruit plants and epiphyte species and to the natural regeneration process within Atlantic forest remnants.     

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.     

To Kill,Stay or Flee: The Effects of Lions and Landscape Factors on Habitat and Kill Site Selection of Cheetahs in South Africa

Understanding how animals utilize available space is important for their conservation, as it provides insight into the ecological needs of the species, including those related to habitat, prey and inter and intraspecific interactions. We used 28 months of radio telemetry data and information from 200 kill locations to assess habitat selection at the 3^rd order (selection of habitats within home ranges) and 4^th order (selection of kill sites within the habitats used) of a reintroduced population of cheetahs Acinonyx jubatus in Phinda Private Game Reserve, South Africa. Along with landscape characteristics, we investigated if lion Panthera leo presence affected habitat selection of cheetahs. Our results indicated that cheetah habitat selection was driven by a trade-off between resource acquisition and lion avoidance, and the balance of this trade-off varied with scale: more open habitats with high prey densities were positively selected within home ranges, whereas more closed habitats with low prey densities were positively selected for kill sites. We also showed that habitat selection, feeding ecology, and avoidance of lions differed depending on the sex and reproductive status of cheetahs. The results highlight the importance of scale when investigating a species’ habitat selection. We conclude that the adaptability of cheetahs, together with the habitat heterogeneity found within Phinda, explained their success in this small fenced reserve. The results provide information for the conservation and management of this threatened species, especially with regards to reintroduction efforts in South Africa.     

Do Lions Panthera leo Actively Select Prey or Do Prey Preferences Simply Reflect Chance Responses via Evolutionary Adaptations to Optimal Foraging?

Research on coursing predators has revealed that actions throughout the predatory behavioral sequence (using encounter rate, hunting rate, and kill rate as proxy measures of decisions) drive observed prey preferences. We tested whether similar actions drive the observed prey preferences of a stalking predator, the African lion Panthera leo. We conducted two 96 hour, continuous follows of lions in Addo Elephant National Park seasonally from December 2003 until November 2005 (16 follows), and compared prey encounter rate with prey abundance, hunt rate with prey encounter rate, and kill rate with prey hunt rate for the major prey species in Addo using Jacobs' electivity index. We found that lions encountered preferred prey species far more frequently than expected based on their abundance, and they hunted these species more frequently than expected based on this higher encounter rate. Lions responded variably to non-preferred and avoided prey species throughout the predatory sequence, although they hunted avoided prey far less frequently than expected based on the number of encounters of them. We conclude that actions of lions throughout the predatory behavioural sequence, but particularly early on, drive the prey preferences that have been documented for this species. Once a hunt is initiated, evolutionary adaptations to the predator-prey interactions drive hunting success.     

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.     

Hiding from the moonlight: luminosity and temperature affect activity of Asian nocturnal primates in a highly seasonal forest

The effect of moonlight and temperature on activity of slow lorises was previously little known and this knowledge might be useful for understanding many aspects of their behavioural ecology, and developing strategies to monitor and protect populations. In this study we aimed to determine if the activity of the pygmy loris (Nycticebus pygmaeus) is affected by ambient temperature and/or moonlight in a mixed deciduous forest. We radio-collared five females and five males in the Seima Protection Forest, Cambodia, in February to May, 2008 and January to March, 2009 and recorded their behaviour at 5 minutes intervals, totalling 2736 observations. We classified each observation as either inactive (sleeping or alert) or active behaviour (travel, feeding, grooming, or others). Moon luminosity (bright/dark) and ambient temperature were recorded for each observation. The response variable, activity, was binary (active or inactive), and a logit link function was used. Ambient temperature alone did not significantly affect mean activity. Although mean activity was significantly affected by moonlight, the interaction between moonlight and temperature was also significant: on bright nights, studied animals were increasingly more active with higher temperature; and on dark nights they were consistently active regardless of temperature. The most plausible explanation is that on bright cold nights the combined risk of being seen and attacked by predators and heat loss outweigh the benefit of active behaviours.     

Habitat use and movements of plains zebra (Equus burchelli) in response to predation danger from lions

Prey species must adapt their behavior to avoid predation. Asa key prey item for lions (Panthera leo), plains zebras (Equusburchelli) were expected to respond to immediate threats posedby lions in their area. In addition, zebras were predicted toexhibit behavior tuned to reduce the potential for encounterswith lions, by modifying their movement patterns in the timesof day and habitats of greatest lion danger. We studied a populationof approximately 600 plains zebra living in Ol Pejeta Conservancy,Kenya. We found that zebra abundance on or near a grasslandpatch was lower if lions had also been observed on that patchduring the same day. Predation danger was highest in grasslandhabitat during the night, when lions were more active. Zebrasightings and global positioning system radio collar data indicatedthat zebras also reduced their use of grassland at night, insteadusing more woodland habitat. Zebras moved faster and took sharperturns in grassland at night. It is hypothesized that these moreerratic movements assist zebras in avoiding detection or captureby lions.     

Movement,activity and hunting behaviour of leopards in the Cederberg mountains,South Africa

Few data are available on the behaviour of leopards in the absence of competing large predators and human impact, both of which are believed to influence leopard activity and movements. Remote camera traps and global positioning system (GPS) collars were used to quantify leopard activity in the Cederberg Mountains, seasonal and sexual differences in their movements, and determine whether nocturnal hunting success was related to lunar activity. Seventy‐seven per cent of camera‐trap photographs were at night, with a strong male bias (69%) in captures. Daily displacement using one location per day suggested that males moved significantly further than females. However, multiple locations (≥6 per day) showed no difference because males moved in a more linear fashion, but not further each day, than females. In the Cederberg Mountains, an open rocky habitat with low human impact and no competing predators, leopards were predominantly nocturnal, mainly hunting diurnal prey species. Hunting success was low: leopards travelled long distances between kills, with nocturnal hunting success higher on darker nights.     

The largest European lion Panthera leo spelaea (Goldfuss 1810) population from the Zoolithen Cave,Germany: specialised cave bear predators of Europe

Remains of 13 individuals with 3/1 male/female ratio of the extinct Upper Pleistocene lion Panthera leo spelaea (Goldfuss, 1810) from the Zoolithen Cave near Burggeilenreuth (Bavaria, Germany) include the holotype skull and all paratype material. The highest mortality rate for the Zoolithen Cave lions is in their reproductive adult ages. Bite marks on lion bones or skulls are results of hyena activities, or rare cannibalism of lions under stress situations. Lions were possibly also killed in battles with cave bears during predation on hibernating bears in winter times. This cave bear hunt specialisation in caves overlaps with the ecological behaviour of cave bear feeding by Ice Age-spotted hyenas. Both largest Ice Age predators, lions and hyenas, had to specialise on feeding herbivorous cave bears in boreal forest mountainous cave rich regions, where the mammoth steppe megafauna prey was absent. This cave bear hunt by felids, and scavenging by hyenas and other large carnivores such as leopards and wolves explains why cave bears hibernated deep in to the European caves, for protection reasons against predators. Within such lion–cave bear and even lion–hyena conflicts in the caves lions must have been killed sometimes, explaining mainly the skeleton occurrences in different European caves.     

Moonlight Makes Owls More Chatty

Background

Lunar cycles seem to affect many of the rhythms, temporal patterns and behaviors of living things on Earth. Ambient light is known to affect visual communication in animals, with the conspicuousness of visual signals being largely determined by the light available for reflection by the sender. Although most previous studies in this context have focused on diurnal light, moonlight should not be neglected from the perspective of visual communication among nocturnal species. We recently discovered that eagle owls Bubo bubo communicate with conspecifics using a patch of white throat plumage that is repeatedly exposed during each call and is only visible during vocal displays.

Methodology/Principal Findings

Here we provide evidence that this species uses moonlight to increase the conspicuousness of this visual signal during call displays. We found that call displays are directly influenced by the amount of moonlight, with silent nights being more frequent during periods with no-moonlight than moonlight. Furthermore, high numbers of calling bouts were more frequent at moonlight. Finally, call posts were located on higher positions on moonlit nights.

Conclusions/Significance

Our results support the idea that moon phase affects the visual signaling behavior of this species, and provide a starting point for examination of this method of communication by nocturnal species.     

Sport Hunting,Predator Control and Conservation of Large Carnivores

Sport hunting has provided important economic incentives for conserving large predators since the early 1970''s, but wildlife managers also face substantial pressure to reduce depredation. Sport hunting is an inherently risky strategy for controlling predators as carnivore populations are difficult to monitor and some species show a propensity for infanticide that is exacerbated by removing adult males. Simulation models predict population declines from even moderate levels of hunting in infanticidal species, and harvest data suggest that African countries and U.S. states with the highest intensity of sport hunting have shown the steepest population declines in African lions and cougars over the past 25 yrs. Similar effects in African leopards may have been masked by mesopredator release owing to declines in sympatric lion populations, whereas there is no evidence of overhunting in non-infanticidal populations of American black bears. Effective conservation of these animals will require new harvest strategies and improved monitoring to counter demands for predator control by livestock producers and local communities.     

Opportunistic feeding by lions: non-preferred prey comprise an important part of lion diets in a habitat where preferred prey are abundant

Optimal foraging theory predicts less diverse predator diets with a greater availability of preferred prey. This narrow diet niche should then be dominated by preferred prey, with implications for predator–prey dynamics and prey population ecology. We investigated lion (Panthera leo) diets in Hluhluwe–iMfolozi Park (HiP), South Africa, to assess whether lions in a site with a high density of preferred prey (prey species weighing 92–632 kg as estimated from a published meta-analysis) have a narrow diet, consisting primarily of preferred prey. HiP is a useful study site to investigate this prediction because it is a productive landscape (with a high density of prey) where lion-preferred prey constitutes up to 33% of the prey available to lions. Furthermore, to investigate whether lions in HiP exhibit sex-specific diets as documented in other southern African populations, we estimated male and female lion diets separately. We were specifically interested in testing whether traditional approaches of estimating lion diets at the population level mask sex-specific predation patterns, with possible implications for management of lions in small to medium-sized fenced reserves. Lions in HiP preferred larger prey species (63–684 kg) and had diets with a larger proportion of preferred prey than reported in an African-wide meta-analysis. However, despite the high density of preferred prey species, 36% of lion diets still consisted of typically non-preferred species such as nyala (Tragelaphus angasii). This finding suggests that lions in HiP maintain a degree of opportunism even when preferred prey are abundant. Therefore, abundant, non-preferred prey are likely to be an important resource for lion populations. Sex-specific differences in lion diets were evident in HiP, suggesting that estimation of lion resource use and carrying capacity should consider opportunistic hunting and sex-specific differences in lion diets.     

Winter field use and habitat selection by Eurasian Golden Plovers Pluvialis apricaria and Northern Lapwings Vanellus vanellus on arable farmland

Eurasian Golden Plovers Pluvialis apricaria and Northern Lapwings Vanellus vanellus winter in large numbers on Britain's farmland. Previous studies in mixed farming areas showed pronounced preferences for permanent pasture, but increasing numbers of plovers winter in eastern Britain where arable farmland dominates. We show that an area of intensive arable farmland supported significant numbers of both species from October to February in the absence of large areas of pasture. Habitat use varied seasonally as the habitats present changed. Throughout winter, use of cereal crops matched their availability: plovers selected harrow in early winter, sugar beet stubbles in midwinter and other crops in late winter. Pastures were rarely used at any time of the year. Flocks occupied only a fraction of the available fields, concentrating most in large fields with open boundaries and where manure had been applied. Daytime feeding was more likely during cold days after nights with a new moon, short duration of moonlight or low-intensity moonlight. These results show that plovers are currently able to utilize intensive arable farmland but future changes in management, such as cessation of manure applications and reductions in sugar beet cropping, could have detrimental effects.