Exploring African lion (<Emphasis Type="Italic">Panthera leo</Emphasis>) behavioural phenotypes: individual differences and correlations between sociality,boldness and behaviour

Increasing our understanding of personality, at an individual and group level, is crucial to the pre-release assessment of social species within ex situ reintroduction programs. We conducted the first exploration into the personality of a captive-origin pride of African lions (Panthera leo), assessing behavioural variations and consistencies in daily activity, social and hunting behaviour, and boldness. Data were collected via direct observations, while a species-specific protocol for testing boldness, using playbacks, was developed. Differences in sex, age and session time for the activity budget were evaluated using Pearson correlations and repeated-measures ANOVA, while social interactions were analysed using social network analysis. Spearman’s correlations were conducted to assess for associations between boldness scores, activity and sociality. The two boldness tests provided a range of scores per lion, indicating that the test was effective. Correlations and variations in individual behaviour indicated that adults and sub-adults have specific roles within pride behaviour. Correlations between boldness and activity and social behaviours provided information on the role of individuals, allowing investigation into the behaviour of a dominant and a social keystone. Our study indicates that evaluating various aspects of behaviour in conjunction with boldness has the potential to assist the pre-release assessment of a pride within an ex situ reintroduction program.     

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.     

A Social Network Analysis of Social Cohesion in a Constructed Pride: Implications for Ex Situ Reintroduction of the African Lion (Panthera leo)

Animal conservation practices include the grouping of captive related and unrelated individuals to form a social structure which is characteristic of that species in the wild. In response to the rapid decline of wild African lion (Panthera leo) populations, an array of conservational strategies have been adopted. Ex situ reintroduction of the African lion requires the construction of socially cohesive pride structures prior to wild release. This pilot study adopted a social network theory approach to quantitatively assess a captive pride’s social structure and the relationships between individuals within them. Group composition (who is present in a group) and social interaction data (social licking, greeting, play) was observed and recorded to assess social cohesion within a released semi-wild pride. UCINET and SOCPROG software was utilised to represent and analyse these social networks. Results indicate that the pride is socially cohesive, does not exhibit random associations, and the role of socially influential keystone individuals is important for maintaining social bondedness within a lion pride. These results are potentially informative for the structure of lion prides, in captivity and in the wild, and could have implications for captive and wild-founder reintroductions.     

Persistence and local extinction of lion prides in the Ngorongoro Crater, Tanzania

African lions (Panthera leo) live in social groups (prides) that exhibit group territorial behavior. Pride persistence is expected to depend on its ability to compete against neighboring prides as well as on average rates of reproduction and survival, thus providing a meaningful measure of intergenerational reproductive success. We used Akaike’s Information Criterion (AIC) to select the best approximating models explaining how demographic variables influenced pride persistence during a 30-year period in the Ngorongoro Crater, Tanzania, and identified landscape factors affecting those demographic variables. Pride persistence to 10 years depended on adult female density (pride size) and cub productivity (the ultimate source of new females). Average age of adult females had a weakly positive influence on pride persistence, while the effect of female mortality was weakly negative. Adult female mortality increased with disease epidemics and in territories with high human disturbance. Cub productivity was highest in territories closest to rivers and only slightly higher near swamps, and also high in areas of higher vegetative cover and high human use. No landscape variable significantly affected female density. The growth and population size of the Crater lions was closely linked to demographic performance of individual prides, while territorial behavior played a key role in mediating the interactive effects of landscape and demography.     

Social factors influencing reproduction in wild lions

The paper summarizes seven years of records of the reproductive history of two wild prides of lions ( Panthera leo L.) in the Serengeti National Park, Tanzania. These records are analyzed to demonstrate ways in which lion reproduction is influenced by their social group and by changes in that group.
Females gave birth at any time of year; they did so sooner if their previous litter died. Females synchronized births with other pride members. Takeover of the pride by new males resulted in a drop in the birth rate. Most cubs died, before two years old. Birth synchrony resulted in better cub survival. The arrival of new males resulted in an increase in cub mortality. Females came into oestrus synchronously with other pride members. Males had a much longer effective reproductive life if they had male companions.
The proximate causes of these features of lion reproduction are considered, and the evolution of aspects of the organization of reproduction in lions is discussed.     

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

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

Changes in home range size of African lions in relation to pride size and prey biomass in a semi‐arid savanna

Within‐population studies are needed to investigate the extent of, and the factors underlying, intraspecific variation in home range size. We used data from 12 female and 8 male adult lions instrumented with GPS radio‐collars to describe the ranging behaviour of lions in a population from a dystrophic semi‐arid savanna, Hwange National Park, Zimbabwe. We measured prey availability at the home range scale in 2003, 2004, and 2005. For females, home range size increased as pride biomass increased, which is strongly suggestive of expansionism. Once controlled for pride biomass, home range size decreased as prey biomass increased. Pride ranges responded to changes in food abundance on an annual timescale rather than on a seasonal timescale. Female home range size was influenced by the abundance of kudu in the early dry season, whereas it was influenced by buffalo and young elephant abundance in the late dry season. This study shows that female home range size is mainly driven by the size of the pride, but also by prey abundance. Furthermore, female seasonal home range size may be determined, not only by prey abundance, but also by prey dispersion in the landscape. Home range size of males was driven by both prey biomass and the density of female prides.     

Ecological correlates of lion social organization (Panthers, leo)

Data on lion home range size, pride size, density, group size, cub survival, litter size and adult sex ratio were extracted from studies carried out in 10 habitats. The data were compared with five measures of food supply: mean prey biomass, prey biomass during the season of greatest abundance, prey biomass during the season of least abundance, biomass of middle-sized prey species (i.e. those with mean adult live-weights between 50 and 250 kg), and the mean size of carcasses fed on by lions. The results indicated that range size is inversely correlated with the abundance of prey during the period of least abundance, but not with overall prey abundance. Range size showed no consistent variation with pride size or with minimum metabolic requirements. Pride size (measured in terms of average number of animals per pride and average number of adult females) and cub survival strongly correlated with lean season food abundance. No relationship was found between group size or litter size and any of the measures of food supply Adult sex ratio did not vary consistently with food supply or lion density, although the data did suggest that prides inhabiting small, circumscribed reserves may experience less inter-male competition and this, in turn, may affect the adult sex ratio.     

Contraception, reproduction and demography of free-ranging Etosha lions (Panthera leo)

The use of long-acting contraceptives as an effective population control mechanism for free-ranging lions was investigated during a three-year study in the Etosha National Park on five lion prides. Lions were immobilized with ketamine hydrochloride and xylazine hydrochloride. Following immobilization, lionesses were weighed, measured and aged, while serum steroid hormone levels were analysed and vaginal smears were obtained. Physical condition and sexual state were assessed. Individuals were used as control animals or given melengestrol acetate (contraceptive) implants, branded and released; the animals were observed for possible changes in behaviour, birth rate and mortality. Ten treated lionesses were recaptured at irregular intervals to reassess weight and steroid hormone levels, while three lionesses had their implants removed to determine if their fertility would return. The contraceptives prevented pregnancy, were reversible when removed and did not alter lion behaviour significantly, except that sexual behaviour was not recorded in treated lionesses.     

Translocating lions into an inbred lion population in the Hluhluwe-iMfolozi Park, South Africa

A fundamental problem in conservation biology is the risk of inbreeding in fragmented and declining populations. In the Hluhluwe-iMfolozi Park (HiP), a small, enclosed reserve in South Africa, a large lion Panthera leo population arose from a founder group of five individuals in the 1960s. The HiP lion population went through a persistent decline and showed indications of inbreeding depression. To restore the genetic variation of the inbred HiP lion population, new lions were translocated into the existing population. Translocated females formed stable associations and established enduring pride areas with other translocated lionesses, but did not bond into native female prides. The translocated male coalition was more successful in gaining and maintaining residence in a pride than the translocated lone male that split off on his own from the male coalition. Litter size and cub survival was about twice as high for pairings involving at least one translocated parent than for pairings of two native lions. It is therefore possible to infuse new genes rapidly and successfully into a small, isolated lion population. Such translocations may become an important adaptive management tool as lion populations become increasingly fragmented.     

Habitat selection and feeding ecology of the cheetah (Acinonyx jubatus) in thicket vegetation: is the cheetah a savanna specialist?

Habitat selection and feeding ecology of a reintroduced population of cheetahs Acinonyx jubatus were studied in a 16 000 ha game reserve in the Eastern Cape Province (South Africa). Seventy per cent of the reserve is characterized by very dense thicket vegetation (valley bushveld) and the remainder is open and savanna-like. The results illustrated a strong effect of sex and group size on the behaviour of cheetahs. The coalition (three adult males) killed significantly larger animals (55% of kills weighed more than 65 kg) than single female cheetahs (less than 2% of kills weighed more than 65 kg). Female cheetahs showed temporal and spatial avoidance of lions by hunting at dawn and dusk and positioning their home ranges [95% utilization distribution (UDs)] significantly farther from the pride of lions than did the coalition. The coalition hunted earlier and later than female cheetahs, and 46% of their kills were made in darkness. In addition, their home range overlapped that of the lions and they showed neither temporal nor spatial avoidance of the lions. The rates of kleptoparasitism were lower and the kill retention times were longer than those reported elsewhere in Africa, and it is suggested that this is a consequence of the cover provided by the thicket vegetation and prey size. The home ranges (95% UDs) of female cheetahs incorporated more thicket vegetation than that of the coalition, indicating that the coalition is less susceptible to predation than single females. These data suggest that cheetahs possess greater behavioural flexibility than previously reported, that they can hunt successfully in thicket vegetation, sometimes in darkness, that they are not restricted to killing small to medium-sized prey, and that they may not be savanna specialists.     

Home Range Utilisation and Territorial Behaviour of Lions (Panthera leo) on Karongwe Game Reserve,South Africa

Interventionist conservation management of territorial large carnivores has increased in recent years, especially in South Africa. Understanding of spatial ecology is an important component of predator conservation and management. Spatial patterns are influenced by many, often interacting, factors making elucidation of key drivers difficult. We had the opportunity to study a simplified system, a single pride of lions (Panthera leo) after reintroduction onto the 85 km² Karongwe Game Reserve, from 1999–2005, using radio-telemetry. In 2002 one male was removed from the paired coalition which had been present for the first three years. A second pride and male were in a fenced reserve adjacent of them to the east. This made it possible to separate social and resource factors in both a coalition and single male scenario, and the driving factors these seem to have on spatial ecology. Male ranging behaviour was not affected by coalition size, being driven more by resource rather than social factors. The females responded to the lions on the adjacent reserve by avoiding the area closest to them, therefore females may be more driven by social factors. Home range size and the resource response to water are important factors to consider when reintroducing lions to a small reserve, and it is hoped that these findings lead to other similar studies which will contribute to sound decisions regarding the management of lions on small reserves.     

Behavioural differences between two captive populations of red jungle fowl (Gallus gallus) with different genetic background,raised under identical conditions

Ex situ conservation of threatened species may lead to behavioural adaptation, which can affect success of reintroduction attempts. In previous studies, we investigated the effects of captivity on the behaviour of red jungle fowl (Gallus gallus) and found that captive populations differed behaviourally as well as genetically. The aim of the present study was to compare the behaviour of two of the previously studied populations, raised under identical conditions. Eggs were collected from birds at Copenhagen zoo (Cop) and Götala research station (Got) and were incubated and hatched together. Twenty-eight birds (16 Got and 12 Cop) were reared together and tested in eight different behavioural tests, measuring different aspects of fear-related behaviours as well as exploratory and social behaviours. The study revealed several differences in fear-related behaviours between the populations but none in exploratory or social behaviours. In general, one of the populations (Cop) showed more intense fear behaviours than the other (Got), which instead were less fearful in their behaviours. This indicates that breeding animals in captivity may lead to behavioural modifications, which can affect the outcome of reintroductions. The results further suggest that fear-related behaviours are dependent on the genetic background of the animals while social behaviours may be more influenced by the social environment. Since fear-related behaviours, such as predator avoidance and fear of humans, are essential for a life in the wild, these aspects are crucial for the breeding of animals in captivity for conservation purposes.     

Subspecific identity and a comparison of genetic diversity between wild and ex situ wildebeest

The original North American ex situ wildebeest population was believed to originate from the white-bearded wildebeest (Connochaetes taurinus albojubatus), which is both morphologically distinct and geographically separated from the brindled wildebeest (C. t. taurinus). However, after an import of wildebeest into North America in 2001, managers have suspected that white-bearded and brindled wildebeest were mixed in herds at multiple institutions. We sequenced the mitochondrial control region (d-loop) from a portion of the managed North American population and compared our sequences with previously published sequences from wild individuals to determine the subspecific identity and genetic diversity of our ex situ population. We were able to confidently identify C. t. albojubatus as the subspecies identity of the sampled portion of our population. Within our population, haplotype and nucleotide diversity were low (0.169 and 0.001, respectively) with a single common haplotype (H1) containing 41 of the 45 individuals sequenced, while two rare haplotypes (H2 and H3) were derived from three individuals and a single individual, respectively. Nucleotide and haplotype diversity were greater overall in the wild populations compared with our managed population. However, C. t. albojubatus was found to exhibit lower nucleotide diversity in both wild and ex situ populations when compared to other wild subspecies. Though the overall goal of the North American wildebeest population is for public education and not reintroduction, maintaining genetic diversity is vital for the long-term viability of this managed population, which may benefit from periodic supplementation of wild animals.     

The Genetic Integrity of the Ex Situ Population of the European Wildcat (Felis silvestris silvestris) Is Seriously Threatened by Introgression from Domestic Cats (Felis silvestris catus)

Studies on the genetic diversity and relatedness of zoo populations are crucial for implementing successful breeding programmes. The European wildcat, Felis s. silvestris, is subject to intensive conservation measures, including captive breeding and reintroduction. We here present the first systematic genetic analysis of the captive population of Felis s. silvestris in comparison with a natural wild population. We used microsatellites and mtDNA sequencing to assess genetic diversity, structure and integrity of the ex situ population. Our results show that the ex situ population of the European wildcat is highly structured and that it has a higher genetic diversity than the studied wild population. Some genetic clusters matched the breeding lines of certain zoos or groups of zoos that often exchanged individuals. Two mitochondrial haplotype groups were detected in the in situ populations, one of which was closely related to the most common haplotype found in domestic cats, suggesting past introgression in the wild. Although native haplotypes were also found in the captive population, the majority (68%) of captive individuals shared a common mtDNA haplotype with the domestic cat (Felis s. catus). Only six captive individuals (7.7%) were assigned as wildcats in the STRUCTURE analysis (at K = 2), two of which had domestic cat mtDNA haplotypes and only two captive individuals were assigned as purebred wildcats by NewHybrids. These results suggest that the high genetic diversity of the captive population has been caused by admixture with domestic cats. Therefore, the captive population cannot be recommended for further breeding and reintroduction.     

Infanticide by Male Lions Hypothesis: A Fallacy Influencing Research into Human Behavior

Recently, familial abuse and killing of children have been correlated with infanticidal behavior by nonhuman male animals which is postulated to be genetic. However, infanticide by males as an evolutionary mechanism is unlikely to occur in primates, and is equally unlikely in lions. To investigate the sexual selection hypothesis of infanticide for lions, four questions should be considered: (1) because they all mate freely with lionesses, are male pride lions closely related? (often they are not), (2) do males kill the young cubs when they join a pride? (few such cases have been observed), (3) do the females then mate with the new males to produce offspring? (yes), and (4) do the males remain with the pride for over two years to protect this new generation? (often not). Field data indicate that in NO instance do we know that male lions killed cubs in a new pride, mated with their mother, and then remained with the pride to protect their own young. Rather, female lions may kill more cubs than males and certainly cause many more cub deaths by allowing them to starve, [ infanticide by male lions, infanticide hypothesis, faulty science]     

麋鹿发情期主要活动的时间分配及行为研究

本文报道麋鹿在白天主要处于休息状态,早晨、黄昏是它们的摄食时间;成年雌体中的各项活动频率未显示出有明显差异;但雄鹿与雌鹿之间在活动时间分配上有明显差异,而且,雄鹿中的统治者与被统治者之间也表现出有所不同。在攻击行为方面,群内表现出有不同的特权形式;而在交往行为中,发现较多的频次出现于相同性别、年龄和同一阶层中。     

Optimizing the genetic management of reintroduction projects: genetic population structure of the captive Northern Bald Ibis population

Many threatened species are bred in captivity for conservation purposes and some of these programmes aim at future reintroduction. The Northern Bald Ibis, Geronticus eremita, is a Critically Endangered bird species, with recently only one population remaining in the wild (Morocco, Souss Massa region). During the last two decades, two breeding programs for reintroduction have been started (in Austria and Spain). As the genetic constitution of the founding population can have strong effects on reintroduction success, we studied the genetic diversity of the two source populations for reintroduction (‘Waldrappteam’ and ‘Proyecto eremita’) as well as the European zoo population (all individuals held ex situ) by genotyping 642 individuals at 15 microsatellite loci. To test the hypothesis that the wild population in Morocco and the extinct wild population in the Middle East belong to different evolutionary significant units, we sequenced two mitochondrial DNA fragments. Our results show that the European zoo population is genetically highly structured, reflecting separate breeding lines. Genetic diversity was highest in the historic samples from the wild eastern population. DNA sequencing revealed only a single substitution distinguishing the wild eastern and wild western population. Contrary to that, the microsatellite analysis showed a clear differentiation between them. This suggests that genetic differentiation between the two populations is recent and does not confirm the existence of two evolutionary significant units. The European zoo population appears to be vital and suitable for reintroduction, but the management of the European zoo population and the two source populations for reintroductions can be optimized to reach a higher level of admixture.     

African Wild Dogs as a Fugitive Species: Playback Experiments Investigate How Wild Dogs Respond to their Major Competitors

It has been suggested that African wild dogs Lycaon pictus need exceptionally large home ranges (and hence occur at such low densities) because they are limited by competition with larger sympatric carnivores, namely lions Panthera leo and spotted hyenas Crocuta crocuta. To investigate this relationship at a proximate level and explore which factors mediate it, we conducted audio playback experiments examining how wild dogs responded to the simulated proximity of either lions or hyenas. The principle finding was that wild dogs consistently moved directly away from lion roars, but when played hyena whoops either stood their ground or, later, moved off in a random direction. These results suggest that lions represent an immediate high‐level threat to wild dogs that is invariably best avoided, whilst the threat from hyenas may not be so great or perhaps is simply unavoidable. Wild dogs appeared to make some assessment of ambush risk during interactions with lions, illustrated by the varying latency to their retreat in habitats of differing vegetation density (and hence ambush potential). Additionally, packs with younger pups were more likely to alarm call and exhibited a slower rate of retreat in the hour following exposure to lion roars. Other variables investigated (competitor group size, lion sex, presence of pups) failed to explain variation in wild dogs’ responses.     

Genetic structure of lions (Panthera leo L.) in the Selous Game Reserve: implications for the evolution of sociality

Abstract We use 14 microsatellites to examine the genetic structure of a lion (Panthera leo L.) population in southern Tanzania. Heterozygosity levels were high (0.75 ± 0.08). Relatedness estimates showed that prides often had close relatives in neighbouring prides, whereas few relatives were found in prides not sharing a border. The drop‐off in relatedness with distance was highly significant. Female pridemates exhibited a higher mean relatedness (0.26 ± 0.07) to one another than did pride males (0.11 ± 0.07). Mean relatedness among females was significantly higher in small prides than in large ones. Prides exhibited significant inbreeding avoidance (F_IL: ?0.11). Mating did not detectably differ from random across prides (F_IT: ?0.02 ns). In addition to being recognizable behavioural and demographic units, prides were statistically distinct genetic units (F_LT: 0.07). Some neighbouring prides grouped together both geographically and genetically, forming ‘superprides’ in the population (F_ZT = 0.05). Thus, although individual prides were genetically distinct, there was an important genetic structure above the level of social groups.