Competitive interactions between spotted hyenas and lions in the Etosha National Park, Namibia

Spotted hyenas are successful hunters, but they also scavenge. Their main food competitors are lions. In the Etosha National Park, Namibia hyenas are unable to prevent kleptoparasitism by lions and fail to acquire kills from lions. The reasons are the small ratio of hyenas to female and subadult lions at kills and the presence of adult male lions. Because of the hyenas’ small clan sizes and large territories they seem to be unable to recruit sufficient clan members to take over lion kills or deter lions from their own kills. In Etosha, 71% of hyena mortality was due to lions; four cubs and one adult female hyena were killed by male lions during a 1‐year study. Hyenas have evolved adaptations against lions and initiate aggressive interactions with lions without the immediate availability of food, which is termed mobbing behaviour. Etosha hyenas initiated mobbing attempts when lions were near the hyena's communal den. Possibly, Etosha hyenas mobbed lions to distract lions from the hyenas’ den and their cubs and to warn their dependent offspring to hide from lions.     

The holotypes of the upper Pleistocene Crocuta crocuta spelaea (Goldfuss, 1823: Hyaenidae) and Panthera leo spelaea (Goldfuss, 1810: Felidae) of the Zoolithen Cave hyena den (South Germany) and their palaeo‐ecological interpretation

The rediscovered holotype skulls of Late Pleistocene Panthera leo spelaea ( Goldfuss, 1810 ) (Felidae) and Crocuta crocuta spelaea ( Goldfuss, 1823 ) (Hyaenidae) from the Zoolithen Cave at Burggeilenreuth, southern Germany, are discussed. The cave became famous mainly due to its rich cave bear bone remains from the late Saalian (OIS 6–8) to Eemian/Weichselian (OIS 3–6) including additionally a third holotype of Ursus spelaeus Rosenmüller, 1794 (Ursidae). The ‘Felis spelaea’ holotype represents an adult male with a strong bite mark on the saggital crest, which was in an early stage of healing. Compared with other European Late Pleistocene lion skulls and skeletons, and with modern African lions, it provides evidence of intraspecific conflict between male Ice Age lions. The holotype of ‘Hyaena spelaea’ is one of several hundred hyena remains from a well‐frequented hyena den cave. The cave was used intensively by Late Pleistocene hyena clans, for collecting lion carcasses in addition to their accustomed prey, as happened in many caves throughout Europe. Ice Age spotted hyena clans might have killed Ice Age steppe lions for many reasons, such as fights over prey and territory, and the protection of cubs, but they did not always scavenge on their carcasses. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154 , 822–831.     

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.     

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.     

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     

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.     

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.     

Upper Pleistocene Panthera leo spelaea (Goldfuss, 1810) remains from the Bilstein Caves (Sauerland Karst) and contribution to the steppe lion taphonomy,palaeobiology and sexual dimorphism

Remains of the steppe lion Panthera leo spelaea (Goldfuss) from historical digs in the Bilstein Caves of Warstein (Sauerland, NW Germany) are described. Their age seems to be from the Early Weichselian periods (Upper Pleistocene). Whereas the Bilstein cave was inhabited by cave bears at that time only a few hyena prey remains, were most likely imported into the cave entrance by hyenas. Bite and crush marks on a few bones of Bison priscus, Bos primigenius, Cervus elaphus, a rhinoceros Coelodonta antiquitatis vertebra and even several chewed cave bear bones prove the hyena presence which is similar to other caves in the Sauerland hyena den cave rich region. Additionally some larger wolves subspecies Canis lupusspelaeus bones were found, but only few Crocuta crocuta spelaea remains are present. After taphonomic comparisons to six other hyena and cave bear den caves of northern Germany, this cave can be classified as a cave bear den, which was briefly used by hyenas only for food storage or commuting or cave bear predation site in one part of the Cave. The lion material refers at least to one young adult lioness, one more adult female and two male lions; therefore, at minimum, the remains of four adult individuals are represented. The absence of juvenile lion material, in contrast to cave bear cub remains in the Bilstein Caves, proves that P. leo spelaea did not use this and all other caves in the region to raise their cubs. The bone material from the Bilstein Caves would prove the same hyena-lion antagonism conflict being recently proven for the Perick Caves, Balve Cave or Martins Cave well. Other situations in caves such as the Keppler Cave and the Bilstein Cave initially show the more complex taphonomic situation of lion remains in European caves, especially in cave bear dens, where they seem to have hunted periodically cave bears, such as it is already proven for hyenas in the Sauerland Karst and other caves of Europe.     

Lens‐anomalies and other ophthalmic findings in a group of closely‐related angola lions (Panthera leo bleyenberghi)

After the diagnosis of bilateral, immature, nuclear, and posterior cortical cataracts in one Angola lioness, and because of the possible implications of the cataracts for a breeding program, complete ophthalmic examinations on a group of related adult Angola lions and their offspring were carried out. Five adult lions, ranging in age from 1.5–5.5 years, and five lion cubs were studied clinically. The examination included slit‐lamp biomicroscopy, indirect ophthalmoscopy, and photography. The eyes of three of the offspring were submitted for histopathologic examination and examined by light microscopy. The most significant findings were cataracts of various stages, which were observed in four adult lions and one male cub. Mild lenticular abnormalities were noted in the histopathologic examination of the lion cubs' eyes. Additional ophthalmic findings, of lesser clinical consequence, were also noted. This breeding program would benefit from further investigation by animal nutritionists and geneticists, and the animals in this group should undergo periodic ophthalmologic examinations. Zoo Biol 0:1–7, 2006. © 2006 Wiley‐Liss, Inc.     

Threat of rapid extermination of the lion (Panthera leo leo) in Waza National Park,Northern Cameroon

Lion populations in West and Central Africa are small and fragmented. In areas where park management is weak, threats will likely facilitate the extinction of the lion. Wildlife management requires knowledge of the population estimate. The population of lions in Waza National Park (Waza NP) was assessed by individual identification of members in the population. The population was assessed to comprise of 14–21 adult individual lions. The age structure was skewed towards adults; cubs comprised 22% of all lions identified while the sex ratio was 1 : 3. Two out of four collared lions were lost to illegal, retaliatory killings within 1 year; and probably two more males and one more female were also killed during this period. The lion population appears to have declined during the last 5 years with six lions dying per year, which is at a much higher rate than observed in the previous decades. Human‐livestock pressure has increased tremendously in this period, resulting to frequent human‐lion conflicts. To ensure the survival of the lion in Waza NP and in the entire region, management needs to intensify efforts to mitigate the pressure from humans and their livestock.     

人工育幼非洲狮、孟加拉虎、豹幼仔的生长曲线拟合比较

<正>生长曲线(Growth curve)是揭示动物生长发育规律的有效手段之一，反映动物发育指标(包括体重、体尺、体表面积等度量值)与时间之间的关系(盛志廉和吴常信，1995)。生长曲线方程大致可以分为三类：第一类是表示报酬递减规律的方程，如指数函数；第二类是描述光滑S型曲线，     

The response of lions and spotted hyaenas to sound playbacks as a technique for estimating population size

Overall, 217 lions (>1 year) Panthera leo (Linnaeus) and 2069 spotted hyaenas (>8 mo) Crocuta crocuta (Erxleben) responded to broadcast vocalizations, call‐ins, during six surveys in the Masai Mara National Reserve, Kenya.
Carnivore response varied seasonally and was significantly higher and more uniform when migratory prey were absent. Variation in wildebeest abundance, by affecting food availability, was the only significant predictor of response. Variability in response with overall prey abundance was higher for lions than hyaenas. Lion response occurred within a radius of 2·5 km, was independent of age and sex but dropped virtually to zero whenever lions possessed a carcass at the time of broadcasting.
An independent total census recorded 547 lions (all ages) in the Reserve, 9·96% of which were nomads. This proportion increased by 7·94% following the influx of migrants and raised lion (>1 year) density from 0·292 to 0·320 lions/km². The call‐in estimate of 0·294±0·009 (95% c.l.) lions/km² practically equalled the total count. The high precision and low bias (−0·224%) of this estimate imply that call‐in surveys are reliable and may be employed to effectively monitor carnivore populations in the long term. Under simple random sampling, covering about 20% of an area would seem adequate to produce reliable density estimates.     

Spatio-temporal niche partitioning between the African lion (<Emphasis Type="Italic">Panthera leo leo</Emphasis>) and spotted hyena (<Emphasis Type="Italic">Crocuta crocuta</Emphasis>) in western African savannas

Large predators in West Africa are threatened with extinction mainly by direct and indirect effects of human activities. Within this context, intraguild competition can limit populations of some species and even play a role in extinction. In this study, we used camera trapping to assess the spatial and temporal patterns of niche partitioning between the African lion Panthera leo leo and the spotted hyena Crocuta crocuta in Pendjari Biosphere Reserve, Benin. We found that these predators are more nocturnal in the hunting zone than in the national park of the biosphere reserve. The temporal overlap between lion and hyena was high in the national park (Pianka overlap index 0.88) and low in the hunting zones (0.39). The spatial overlap was low (0.40 in the national park and 0.38 in the hunting zones). The two predators were distributed independently in the national park, but showed significant positive association (co-occurrence) in the hunting zones. We suggest that anthropogenic activities leading to depletion of predators and their prey limit lion and hyena distribution in the hunting zones to some safety areas which are strongly selected by both predators. We recommend to significantly improve conservation efforts in the hunting zones of Pendjari Biosphere Reserve and to expand research of lion-hyena intraguild relationships to improve predator survival in West Africa.     

Hunting Patterns,Ban on Baiting,and Harvest Demographics of Brown Bears in Sweden

Abstract We analyzed harvest data to describe hunting patterns and harvest demography of brown bears (Ursus arctos) killed in 3 geographic regions in Sweden during 1981–2004. In addition, we investigated the effects of a ban on baiting, instituted in 2001, and 2 major changes in the quota system: a switch to sex-specific quotas in 1992 and a return to total quotas in 1999. Brown bears (n=887) were harvested specifically by bear hunters and incidentally by moose (Alces alces) hunters. Both hunter categories harvested bears 1) using dogs (37%), 2) by still hunting (30%), 3) with the use of bait (18%), and 4) by stalking (16%). The proportion of bears killed with different harvest methods varied among regions and between bear- and moose-oriented hunters. We found differences between male (52%) and female bears (48%) with respect to the variables that explained age. Moose-oriented hunters using still hunting harvested the youngest male bears. Bears harvested during the first management period (1981–1991) were older and had greater odds of being male than during the subsequent period. It appears that hunters harvesting bears in Sweden are less selective than their North American counterparts, possibly due to differences in the hunting system. When comparing the 4 years immediately prior to the ban on baiting with the 4 years following the ban, we found no differences in average age of harvested bears, sex ratio, or proportion of bears killed with stalking, still hunting, and hunting with dogs, suggesting that the ban on baiting in Sweden had no immediate effect on patterns of brown bear harvest demography and remaining hunting methods. As the demographic and evolutionary side effects of selective harvesting receive growing attention, wildlife managers should be aware that differences in harvest systems between jurisdictions may cause qualitative and quantitative differences in harvest biases. (JOURNAL OF WILDLIFE MANAGEMENT 72(1):79–88; 2008)     

Ultrasonographic and laparoscopic evaluation of the reproductive tract of the captive female African lion (Panthera leo)

The use of transabdominal ultrasonography to assess the oestrous cycle has not been previously described in the African lion (Panthera leo). Twelve sexually mature lionesses and five female cubs had their reproductive organs assessed by transabdominal ultrasound. Ovarian findings were compared to laparoscopic findings while performing laparoscopic ovariectomy or salpingectomy. Vaginal cytology was performed and serum progesterone levels were determined. By combining all data the oestrous cycle stage of each lion was determined. One lion was far pregnant and was not operated on. In adults a uterine body could be seen ultrasonographically in 67% of lions while mural structures could be distinguished in 44% of lions. Five uterine horns could be seen in 3 lions. In 12 adults 10 ovaries were found of which eight had discernable follicles or luteal structures. During laparoscopy 12 active ovaries were seen with luteal structures seen in 11 ovaries and follicles in 2 ovaries. Using laparoscopy as the gold standard, ultrasonography had a sensitivity of 66% and specificity of 83% to detect ovarian reproductive activity. Two uterine cysts and a cluster of periovarian cysts were seen in three different lions. Three lions were pregnant, two were in oestrus, three in a luteal phase (dioestrus), and four were in anoestrus. Transabdominal ultrasound in combination with serum progesterone levels and vaginal cytology can be used to assess ovarian cyclical activity with reasonable accuracy in captive bred lions.     

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.     

Demography of lions in relation to prey and habitat in the Maasai Mara National Reserve, Kenya

We studied lion demography in the Maasai Mara National Reserve between September 1990 and April 1992, with a special emphasis on the spatial and seasonal variation in demographic characteristics. Lion density (0.2–0.4 lions km^?2) and pride size (range 8–48) were high because of a high resident prey biomass (10 335 kg km^?2) augmented by migrant prey to 26 092 kg km^?2 in the dry season. Overall, their sex ratio was almost at parity and varied neither spatially nor seasonally. Sex ratio was even among subadults but skewed toward males and females among cubs and adults, respectively. This implies an increasing differential mortality of males with age through subadulthood. The age ratio varied seasonally because of a birth peak in March–June and an influx of subadults into the reserve during July–August, coincident with increases in migrant prey. The birth peak was apparently preceded by another peak in mating activity falling between November and May. Further research should investigate the precise causes of the biased cub sex ratio, low lion density in the Mara Triangle and the higher ratio of subadults in Musiara than in the Mara Triangle or Sekenani.     

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.     

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.     

Optimal hunting group size: the need for lions to defend their kills against loss to spotted hyaenas

This paper presents preliminary evidence that kleptoparasitism by spotted hyaenas may influence optimal hunting group size of lions. In the absence of adult male lions, hyaenas can drive female and subadult lions off their kills provided they outnumber the lions by a factor of four. Hence the larger the group of lions on the kill, the greater their chance of defending their food against invading hyaenas. At Savuti, where there was a shortage of adult male lions, the groups of female and subadult lions lost almost 20% of their food to hyaenas. Losses were most frequent for those living in small groups. These lions were often satiated by the time the hyaenas acquired the kill, so the hyaenas did not cause immediate need, yet constituted a constant energy drain on lions by forcing them to hunt more frequently. The implications of these observations for modelling optimal hunting group size of lions are discussed.