Eavesdropping and Risk Assessment Between Lions,Spotted Hyenas and African Wild Dogs

Vocalizing allows rapid transmission of detailed information beyond line of sight. However, the risk of eavesdropping by unintended receivers means there is also a potential cost to any vocalization. For fugitive species such as African wild dogs the potential cost of attracting dangerous competitors as eavesdroppers is especially significant. Experiments presented here demonstrate that eavesdropping lions Panthera leo were highly motivated to approach playbacks of wild dog Lycaon pictus vocalizations. As lions will kill any wild dogs they can catch, wild dogs risk paying high costs should their calls be detected. Lions were less likely to approach playbacks of spotted hyena Crocuta crocuta whoops, with responses split according to gender: male lions remained quick to approach hyena whoops, but females without accompanying males typically did not approach. Although hyenas seemed at least as capable as lions of detecting playbacks of wild dog calls, they were significantly less likely to subsequently approach them. Analogous to female lions faced with hyenas, the reluctance of hyenas to approach wild dogs may well derive from an assessment of the potential risks involved. We consider the hypothesis that wild dog twitters display counter‐adaptations against eavesdropping, but suggest that this species may best limit the risk of detection by avoiding areas where they are most likely to be overheard by lions.     

Spatial and temporal avoidance of risk within a large carnivore guild

Within a large carnivore guild, subordinate competitors (African wild dog, Lycaon pictus, and cheetah, Acinonyx jubatus) might reduce the limiting effects of dominant competitors (lion, Panthera leo, and spotted hyena, Crocuta crocuta) by avoiding them in space, in time, or through patterns of prey selection. Understanding how these competitors cope with one other can inform strategies for their conservation. We tested how mechanisms of niche partitioning promote coexistence by quantifying patterns of prey selection and the use of space and time by all members of the large carnivore guild within Liuwa Plain National Park in western Zambia. Lions and hyenas specialized on wildebeest, whereas wild dogs and cheetahs selected broader diets including smaller and less abundant prey. Spatially, cheetahs showed no detectable avoidance of areas heavily used by dominant competitors, but wild dogs avoided areas heavily used by lions. Temporally, the proportion of kills by lions and hyenas did not detectably differ across four time periods (day, crepuscular, early night, and late night), but wild dogs and especially cheetahs concentrated on time windows that avoided nighttime hunting by lions and hyenas. Our results provide new insight into the conditions under which partitioning may not allow for coexistence for one subordinate species, the African wild dog, while it does for cheetah. Because of differences in responses to dominant competitors, African wild dogs may be more prone to competitive exclusion (local extirpation), particularly in open, uniform ecosystems with simple (often wildebeest dominated) prey communities, where spatial avoidance is difficult.     

Responses of Spotted Hyenas to Lions Reflect Individual Differences in Behavior

Competition among mammalian carnivores can be particularly intense and can influence population dynamics at lower trophic levels. One strategy employed by carnivores to minimize potentially costly interspecific competition is avoidance of dominant species. Recent research has highlighted the importance of consistent individual differences in behavior (i.e. temperament traits) in understanding behavioral variation during predator–prey interactions and intraspecific interactions. However, the importance of such individual differences to interspecific competition has received little attention. Here, we examined the responses of spotted hyenas (Crocuta crocuta) to their primary competitors, African lions (Panthera leo), to (1) determine whether hyenas avoid lions and (2) evaluate the potential importance of individual differences in behavior during interspecific competition. Spotted hyenas and lions co‐occur throughout much of Africa and are vigorous competitors. Whereas lions sometimes kill hyenas and steal their food, lions also represent a source of food for hyenas via scavenging. Using audio playback experiments, we found that hyenas do not uniformly avoid potential encounters with lions. Indeed, we noted considerable variation among individuals in their responses to lion roars, and this variation reflected consistent individual differences in risk‐taking and vigilance tendencies. Individual differences in vigilance behavior were specific to interactions with lions. We conclude that individual differences in behavior have the potential to play an important role in determining the nature and outcome of interspecific competition.     

Living with the enemy: avoidance of hyenas and lions by cheetahs in the Serengeti

Predator avoidance is likely to play a strong role in structuringspecies communities, even where actual mortality due to predationis low. In such systems, mortality may be low because predatoravoidance is effective, and if the threat of predation is liftedthen entire community structures may be altered. Where competitionis intense, then competitor avoidance may have a similar impacton communities. Avoidance behaviors have been documented fora wide range of species, but this is the first attempt to documentavoidance behavior within a large carnivore community. Audioplayback techniques are used to examine the risk perceivedby cheetahs from their two main competitors that are also theirmain predators, lions and hyenas. The results from these experimentsshow that cheetahs actively moved away from lion and hyenaplayback experiments, compared with dummy playbacks where no sound was played. Cheetahs showed no differences in their responsesto playbacks dependent on their sex or reproductive status,suggesting they were responding principally to a competitionrather than a predation threat. However, cheetahs were muchless likely to hunt after competitor playbacks than after dummyplaybacks, and this resulted in a lower kill rate after competitorplaybacks, demonstrating that the perceived presence of competitors had a noticeable impact on the foraging rate of cheetahs. Furthermore,while cheetahs moved just as far following lion playbacks asafter hyena playbacks, they spent significantly more time lookingat the loudspeaker and were less likely to make a kill afterlion playbacks, suggesting that cheetahs perceive lions tobe a greater threat than hyenas.     

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.     

Diet quality in a wild grazer declines under the threat of an ambush predator

Predators influence prey populations not only through predation itself, but also indirectly through prompting changes in prey behaviour. The behavioural adjustments of prey to predation risk may carry nutritional costs, but this has seldom been studied in the wild in large mammals. Here, we studied the effects of an ambush predator, the African lion (Panthera leo), on the diet quality of plains zebras (Equus quagga) in Hwange National Park, Zimbabwe. We combined information on movements of both prey and predators, using GPS data, and measurements of faecal crude protein, an index of diet quality in the prey. Zebras which had been in close proximity to lions had a lower quality diet, showing that adjustments in behaviour when lions are within short distance carry nutritional costs. The ultimate fitness cost will depend on the frequency of predator–prey encounters and on whether bottom-up or top-down forces are more important in the prey population. Our finding is the first attempt to our knowledge to assess nutritionally mediated risk effects in a large mammalian prey species under the threat of an ambush predator, and brings support to the hypothesis that the behavioural effects of predation induce important risk effects on prey populations.     

Characteristics of African wild dog natal dens selected under different interspecific predation pressures

To successfully reproduce, many carnivorous mammals need access to suitable den sites. Den site selection is often based on fitness related criteria like escape from predators, food availability and shelter from extreme weather conditions. African wild dogs are cooperative breeders that use a den to give birth to their offspring. They often co-exist with lions and spotted hyenas, both of which are known to kill African wild dog pups. Little is known about den site selection by African wild dogs. In this study, we compared vegetation characteristics and distribution of roads and waterholes around den sites and random sites, in areas with high and low lion and spotted hyena densities. In both areas, African wild dogs selected den sites in closed woodland with little visibility, which is likely to reduce detection by predators, increase the likelihood of escape when detected, and might provide shelter from extreme weather conditions. In the high predator density area, African wild dogs seemed to spatially avoid predators by selecting den sites in this type of habitat relatively further away from waterholes and roads. African wild dogs have high energetic costs of gestation. Therefore, even when predation risk is relatively low, they are likely to try to maximise their fitness by choosing a den site in habitat that will provide optimal protection for their offspring, leaving little additional options to respond to a higher predation pressure.     

A diseased Panthera leo spelaea (Goldfuss 1810) lioness from a forest elephant graveyard in the Late Pleistocene (Eemian) interglacial lake at Neumark-Nord,central Germany

A solitary articulated skeleton of a middle-aged and diseased Panthera leo spelaea lioness from the Eemian interglacial has been found amongst numerous articulated skeletons of Palaeoloxodon antiquus forest elephants, in sediments from a small, shallow lake at Neumark-Nord in central Germany, which has Neanderthal settlements along its shoreline. Several pathologies such as a fibula fracture, arthritis in one of the front legs and a lost canine tooth with associated maxillary inflammation and dissolution made the lioness vulnerable to other predators such as hyenas, whose presence is indicated by their bones, coprolites and many scavenging marks on the elephant skeletons and on a femur from a male lion. The scavenging of hyenas and lions at this site is commonly documented by canine bite marks on the joints of elephant bones. Bite and scratch marks on the ventral vertebral columns and pelvises of two P. antiquus forest elephant skeletons suggest that the intestines and inner organs may have been consumed by large predators, as is commonly the case with modern African lions feeding on elephants. The weak and diseased lioness may possibly have been killed during antagonistic battles between hyenas and lions over their larger prey.     

A survey for Echinococcus spp. of carnivores in six wildlife conservation areas in Kenya

To investigate the presence of Echinococcus spp. in wild mammals of Kenya, 832 faecal samples from wild carnivores (lions, leopards, spotted hyenas, wild dogs and silver-backed jackals) were collected in six different conservation areas of Kenya (Meru, Nairobi, Tsavo West and Tsavo East National Parks, Samburu and Maasai Mara National Reserves). Taeniid eggs were found in 120 samples (14.4%). In total, 1160 eggs were isolated and further analysed using RFLP-PCR of the nad1 gene and sequencing. 38 of these samples contained eggs of Echinococcus spp., which were identified as either Echinococcus felidis (n = 27) or Echinococcus granulosus sensu stricto (n = 12); one sample contained eggs from both taxa. E. felidis was found in faeces from lions (n = 20) and hyenas (n = 5) while E. granulosus in faeces from lions (n = 8), leopards (n = 1) and hyenas (n = 3). The host species for two samples containing E. felidis could not be identified with certainty. As the majority of isolated eggs could not be analysed with the methods used (no amplification), we do not attempt to give estimates of faecal prevalences. Both taxa of Echinococcus were found in all conservation areas except Meru (only E. felidis) and Tsavo West (only E. granulosus). Host species identification for environmental faecal samples, based on field signs, was found to be unreliable. All samples with taeniid eggs were subjected to a confirmatory host species RLFP-PCR of the cytochrome B gene. 60% had been correctly identified in the field. Frequently, hyena faeces were mistaken for lion and vice versa, and none of the samples from jackals and wild dogs could be confirmed in the tested sub-sample. This is the first molecular study on the distribution of Echinococcus spp. in Kenyan wildlife. The presence of E. felidis is confirmed for lions and newly reported for spotted hyenas. Lions and hyenas are newly recognized hosts for E. granulosus s.s., while the role of leopards remains uncertain. These data provide the basis for further studies on the lifecycles and the possible link between wild and domestic cycles of cystic echinococcosis in eastern Africa.     

Lion,Ungulate, and Visitor Reactions to Playbacks of Lion Roars at Zoo Atlanta

Felids in captivity are often inactive and elusive in zoos, leading to a frustrating visitor experience. Eight roars were recorded from an adult male lion and played back over speakers as auditory enrichment to benefit the lions while simultaneously enhancing the zoo visitor experience. In addition, ungulates in an adjacent exhibit were observed to ensure that the novel location and increased frequency of roars did not lead to a stress or fear response. The male lion in this study roared more in the playback phase than in the baseline phases while not increasing any behaviors that would indicate compromised welfare. In addition, zoo visitors remained at the lion exhibit longer during playback. The nearby ungulates never exhibited any reactions stronger than orienting to playbacks, identical to their reactions to live roars. Therefore, naturalistic playbacks of lion roars are a potential form of auditory enrichment that leads to more instances of live lion roars and enhances the visitor experience without increasing the stress levels of nearby ungulates or the lion themselves, who might interpret the roar as that of an intruder.     

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.     

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.     

Urgent call for further breeding of the relic zoo population of the critically endangered Barbary lion (Panthera leo leo Linnaeus 1758)

The Barbary lion became extinct in the wild around 1942. Since the end of the 19th century, a last purebred captive breeding stock existed at the court of Morocco. The rest of these animals became the core exhibition of the new Rabat Zoo after passing through repeated bottlenecks and possibly some introgression events by foreign lions. This study uses mitochondrial DNA sequencing data to clarify the relationship among these lions and their sub-Saharan and Asian relatives. We analysed mitochondrial cytochrome b sequences obtained from a sample from a Barbary lion descended from a young female of the Barbary lion breeding group at the Rabat Zoo and various other members of the genus Panthera. In our cytochrome-b-based phylogenetic tree, the North African Barbary lion, represented by a biopsy sample from the Neuwied Zoo, joins the Asian lion clade, although it is slightly different from its Asian sister group. However, it is clearly distinct from sub-Saharan lions and can be considered as a genetically defined phylogeographic group of its own. Molecular dating of the extant sub-Saharan and Asian lion groups shows that the split between North African Barbary lions and Asian lions must be considerably more recent than 74–203 kilo years ago.     

Livestock Predation—Insights From Problem Animal Control Registers in Botswana

Abstract: Problem Animal Control Registers, where farmers report livestock losses due to predators as a prerequisite for financial compensation, allow quantifying the human-predator conflict. We analyzed such registers from the Kweneng District of Botswana to assess the impact of native predators on livestock over 3 years. Leopards (Panthera pardus), lions (Panthera leo), wild dogs (Lycaon pictus), brown hyenas (Hyaena brunnea), and cheetahs (Acinonyx jubatus) altogether claimed 2,272 head of livestock. During 2002, the year with the highest impact, the number of animals reported depredated (954) represented 0.34% of the livestock in the district. Leopards and lions caused 64% of the losses. Leopard livestock predation mainly affected calves and was consistent over the district and over time. In contrast, lion predation concentrated on adult cattle, was characterized by local hot spots close to reserve borders, decreased with increasing distance to a reserve, and increased during 2002, an unusually dry year. Interviews with 60 farmers and herders within 30 km of Khutse and Central Kalahari Game Reserves revealed an annual loss of 2.2% of their livestock to predators. Here, small farms (max. 100 domestic animals) suffered relatively higher losses than large, commercial farms, not only due to predation (small farms: 11.7%; large farms: 1.0%) but also from other causes (small: 12.6%; large: 2.8%), even though herders on large farms guarded 5 times more livestock per person than those on small farms. To reduce livestock predation in most of the district where lions are absent, we recommend maternity corrals for pregnant females and calves to better protect vulnerable calves during day and night. In areas close to a reserve where lions roam, herders' incentives to keep all livestock protected in a corral at night have to be enhanced because, according to the registers, only 3 predation cases were reported to have happened inside a corral.     

Effects of research disturbance on the behavior and abundance of Steller sea lions (Eumetopias jubatus) at two rookeries in Alaska

We examined the effects of research disturbance on the behavior and abundance of Steller sea lions (Eumetopias jubatus) at rookeries on Marmot and Ugamak Islands in Alaska. During 3 of 6 yr, researchers intentionally drove all adult and juvenile sea lions off at least part of the beach in order to permanently mark and measure sea lion pups. The research disturbance occurred after the majority of females had bred and when most pups were 1 mo old. We used generalized linear models to determine the relationship between research disturbance and sea lion behavior or abundance. Research disturbance was related to changes in the proportion of sea lions exhibiting two to three of nine behavior metrics: agonistic and resting females and active males at Marmot, and active and resting males and females at Ugamak. Model results indicated that changes lasted between 3 and 20 d depending on the sex, behavior, and rookery. Inclusion of research disturbance into Marmot abundance models did not improve the fit to the data, if variability between years was permitted. Optimally timed, low‐frequency research disturbance did not appear to have long‐term effects on sea lion behavior or abundance and was largely associated with changes that were similar to natural variation.     

STELLER SEA LION STATUS AND TREND IN SOUTHEAST ALASKA: 1979–1997

Steller sea lion (Eumetopias jubatus) numbers in the United States declined by about 75% over the past 20+ yr. They are classified, under the U. S. Endangered Species Act, as “threatened” in the eastern portion of their range and as “endangered” in the western portion. We analyzed trends in numbers of pup and non-pup Steller sea lions counted in Southeast Alaska between 1979 and 1997. Sea lion numbers, based on counts of pups on rookeries, increased by an average of 5.9% per year between 1979 and 1997. However, numbers of pups increased at a much slower rate (+ 1.7% per year) between 1989 and 1997. For counts of non-pup Steller sea lions we used models that controlled for the effects of date, time, and tide at the time of the survey to analyze trends. This technique reduced bias and increased precision of the resulting trend estimates. Numbers of sea lions were stable (+0.5%) between 1989 and 1996, based on counts of non-pups. We estimated the Southeast Alaska breeding population of Steller sea lions at about 19,000 animals of all ages in 1997, a level that is probably near the highest in recorded history.     

Rabies Virus and Canine Distemper Virus in Wild and Domestic Carnivores in Northern Kenya: Are Domestic Dogs the Reservoir?

Rabies virus (RV) and canine distemper virus (CDV) can cause significant mortality in wild carnivore populations, and RV threatens human lives. We investigated serological patterns of exposure to CDV and RV in domestic dogs (Canis familiaris), African wild dogs (Lycaon pictus), black-backed jackals (Canis mesomelas), spotted hyenas (Crocuta crocuta), striped hyenas (Hyaena hyaena) and African lions (Panthera leo), over a 10-year period, in a Kenyan rangeland to assess the role domestic dogs may play in the transmission dynamics of these two important canid pathogens. Observed patterns of RV exposure suggested that repeated introduction, rather than maintenance, occurred in the wild carnivore species studied. However, RV appeared to have been maintained in domestic dogs: exposure was more likely in domestic dogs than in the wild carnivores; was detected consistently over time without variation among years; and was detected in juveniles (≤1-year-old) as well as adults (>1-year-old). We conclude that this domestic dog population could be a RV reservoir. By contrast, the absence of evidence of CDV exposure for each carnivore species examined in the study area, for specific years, suggested repeated introduction, rather than maintenance, and that CDV may require a larger reservoir population than RV. This reservoir could be a larger domestic dog population; another wildlife species; or a “metareservoir” consisting of multiple interconnected carnivore populations. Our findings suggest that RV risks to people and wild carnivores might be controlled by domestic dog vaccination, but that CDV control, if required, would need to target the species of concern.     

Evidence for African origins of founders of the asiatic lion species survival plan

The Asiatic lion (Panthera leo persica) exists in the wild as a single relict population of approximately 250 individuals in the protected Gir Forest Sanctuary in western India. In 1981, a species survival plan (SSP) for the Asiatic lion was established by the American Association of Zoological Parks and Aquariums to manage the 200 + descendants of Asiatic lions in captivity in western zoological facilities. This captive population was derived from seven founders. In order to compare the genetic structure of the Gir Forest population with that of the captive SSP population, a genetic survey of 46 electrophoretic allozyme systems resolved from extracts of lion blood was undertaken by using 29 SSP Asiatic lions and 28 wild-caught or captive-bred lions maintained at the Sakkarbaug Zoo in India but originally derived from the Gir Forest. The Gir lion population was found to be genetically monomorphic at each of 46 allozyme loci. This was in contrast to several African lion (Panthera leo leo) populations, which show moderate levels of allozyme variation at the same loci. The SSP lion population was polymorphic at three allozyme loci (IDHI, TF, and PTI) for alleles that were previously found only in African lion populations. Pedigree analysis of the genetic transmission of these three biochemical loci demonstrated that two of the five primary founder animals of the SSP Asiatic lion population (a breeding pair originally imported from the Trivandrum Zoo in southern India) were descendants of the African subspecies. Three other founder animals were pure Asian. A retrospective SSP pedigree analysis of two morphologic characters (prominent abdominal fold and pairing of infraorbital foramen) that are partially diagnostic for persica vs leo was consistent with this conclusion as well. The implications for the management of small captive populations of threatened species and of the Asiatic lion SSP population are discussed.     

A genetically distinct lion (Panthera leo) population from Ethiopia

Lion (Panthera leo) numbers are in serious decline and two of only a handful of evolutionary significant units have already become extinct in the wild. However, there is continued debate about the genetic distinctiveness of different lion populations, a discussion delaying the initiation of conservation actions for endangered populations. Some lions from Ethiopia are phenotypically distinct from other extant lions in that the males possess an extensive dark mane. In this study, we investigated the microsatellite variation over ten loci in 15 lions from Addis Ababa Zoo in Ethiopia. A comparison with six wild lion populations identifies the Addis Ababa lions as being not only phenotypically but also genetically distinct from other lions. In addition, a comparison of the mitochondrial cytochrome b (CytB) gene sequence of these lions to sequences of wild lions of different origins supports the notion of their genetic uniqueness. Our examination of the genetic diversity of this captive lion population shows little effect of inbreeding. Immediate conservation actions, including a captive breeding programme designed to conserve genetic diversity and maintain the lineage, are urgently needed to preserve this unique lion population.     

On the distinctiveness of the Cape lion (Panthera leo melanochaita Smith, 1842), and a possible new specimen from the Zoological Museum, Copenhagen

The Cape lion was a population of lions that probably inhabited the western part of the Cape Province of South Africa until their extermination by man in the mid-19th century. Only a few skeletal remains are known, making every specimen valuable. In this paper, I report on a possible new male specimen CN1570 from the Zoological Museum in Copenhagen. A multivariate discriminant analysis on 27 craniodental variables provided clear separation between four of the five lion subspecies (Panthera leo krugeri, nubica, persica, senegalensis) that were included, whereas P. l. bleyenberghi showed some overlap with P. l. senegalensis and P. l. krugeri. The only two undisputed Cape lion males grouped separately from all other lions. CN1570 also grouped separately from other lions, and towards the two Cape lions. The external morphology of the Cape lion is often cited as having been different from other sub-Saharan lions, but phenotypic plasticity argues for caution in placing emphasis on mane morphology as a distinguishing character among lion subspecies. Skull morphology, however, appears to clearly distinguish male Cape lions from other African lions.