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Factors affecting the hunting success of male and female lions in the Kruger National Park
Abstract
Long-term continuous observations of hunting lions Panthera leo in the Kruger National Park were used to assess the variables affecting hunting success of male and female lions. Generalized linear models revealed that seven variables had significant independent influences on hunting success, with the most important being the prey species hunted. Three types of variables were recognized: (1) lion related, where type of hunt, wind orientation, and the number of adults hunting; (2) prey related, where prey species and herd size; (3) environment related, where moon brightness, and grass height were significant. The sex of the lions had no effect on the overall probability of hunting success. Five second-order interactions significantly influenced the probability of hunting success, with the most important being the interaction between sex and the type of prey. The only significant third-order interaction containing the variable sex, was the inter-relationship with prey species and shrub cover. After removing the over-riding bias of the prey species, greater resolution of the factors affecting success was revealed. The major difference was that group size influences hunting success and concomitantly prey selection, promoting selection for medium-sized ungulates like zebra Equus burchelli and wildebeest Connochaetes taurinus by females, and males mainly capturing buffalo Syncerus caffer. We conclude that in African ecosystems, the hunting success of male and female lions varies with a range of combinations of lion-, prey- and environment-related variables. We demonstrate the hunting ability of male lions, which has perhaps been understated in other studies.
... This trade-off is dependent on the predators' hunting mode, landscape cover and prey behaviour [15][16][17] . For example, stalk and ambush predators, which largely depend on some form of concealment to approach their prey 14,16 , may use darkness as a substitute for physical cover 18 and shift from diurnal to nocturnal hunting as habitats vary from closed to open, despite the main activity of their prey being diurnal [19][20][21] . Under such conditions, the effectiveness of darkness for concealment may decrease in relation to moon luminosity 18,22 , with predator hunting success varying with the moon phase 23 . ...
... For example, stalk and ambush predators, which largely depend on some form of concealment to approach their prey 14,16 , may use darkness as a substitute for physical cover 18 and shift from diurnal to nocturnal hunting as habitats vary from closed to open, despite the main activity of their prey being diurnal [19][20][21] . Under such conditions, the effectiveness of darkness for concealment may decrease in relation to moon luminosity 18,22 , with predator hunting success varying with the moon phase 23 . ...
... Ibex estimates were from hourly data and were therefore rescaled to be made comparable to the 5-hourly snow leopard estimates. . Our data suggest that 'prey catchability' is a more plausible explanation for higher snow leopard activity during the night in all seasons, as, in an open landscape of high visibility above the tree line, these ambush predators would be better able to approach their prey under the cover of darkness and improve their hunting success 18 . However, this does not explain why peak snow leopard activity often occurred during twilight periods rather than in night-time darkness. ...
The daily and seasonal activity patterns of snow leopards (Panthera uncia) are poorly understood, limiting our ecological understanding and hampering our ability to mitigate threats such as climate change and retaliatory killing in response to livestock predation. We fitted GPS-collars with activity loggers to snow leopards, Siberian ibex (Capra sibirica: their main prey), and domestic goats (Capra hircus: common livestock prey) in Mongolia between 2009 and 2020. Snow leopards were facultatively nocturnal with season-specific crepuscular activity peaks: seasonal activity shifted towards night-sunrise during summer, and day-sunset in winter. Snow leopard activity was in contrast to their prey, which were consistently diurnal. We interpret these results in relation to: (1) darkness as concealment for snow leopards when stalking in an open landscape (nocturnal activity), (2) low-intermediate light preferred for predatory ambush in steep rocky terrain (dawn and dusk activity), and (3) seasonal activity adjustments to facilitate thermoregulation in an extreme environment. These patterns suggest that to minimise human-wildlife conflict, livestock should be corralled at night and dawn in summer, and dusk in winter. It is likely that climate change will intensify seasonal effects on the snow leopard's daily temporal niche for thermoregulation in the future.
... The time of day and the lunar phase are also cited as determining factors that influence the feeding behaviour of these animals [46,47]. The species, group size and anti-predatory strategies of the prey, as well as the size and composition of the group and the hunting techniques employed by the «cat herd or pride» (e.g., Lion), will ultimately determine the success or failure of the hunting activities [48,49]. ...
... This has been suggested to enable predators to approach prey of progressively considerably higher sizes with time [70][71][72]. By contrast, no significant differences were reported when the sex of the predator was considered, contrary to other studies reporting a relatively greater probability of success in males than females in some feline predators, such as lions [48]. ...
... Our results suggest the time elapsed between potential prey identification and the first physical contact between predator and prey doubled when prey managed to escape. In this regard, Funston et al. [48] suggests predators that initiate the pursuit of their prey immediately after detection, have a greater probability of success in their capture than those that execute a previous stalking, of variable duration, to the pursuit per se. ...
The individuals engaged in predation interactions modify their adaptation strategies to improve their efficiency to reach success in the fight for survival. This success is linked to either capturing prey (predator) or escaping (prey). Based on the graphic material available on digital platforms both of public and private access, this research aimed to evaluate the influence of those animal- and environment-dependent factors affecting the probability of successful escape of prey species in case of attack by big cats. Bayesian predictive analysis was performed to evaluate the outcomes derived from such factor combinations on the probability of successful escape. Predator species, age, status at the end of the hunting act, time lapse between first attention towards potential prey and first physical contact, prey species and the relief of the terrain, significantly conditioned (p < 0.05) escape success. Social cooperation in hunting may be more important in certain settings and for certain prey species than others. The most parsimonious model explained 36.5% of the variability in escaping success. These results can be useful to design translatable selective strategies not only seeking to boost predation abilities of domestic felids for pest control, but also, biological antipredator defence in potential domestic prey of big cats.
... We note that factors such as prey age-sex class, predator hunting-group size, visibility, time of day, and other social and environmental factors may influence predator capture success, style, and preference and that behaviors such as hunting style lie along a spectrum [34,35,[40][41][42]. We rely on expert knowledge and published literature to classify predator behaviors and consider the following values to represent relative indices or general trends of behavior rather than absolute metrics. ...
... Capture success was defined as the percentage of initiated hunts that ended in a kill. Specific estimates for each predator-prey dyad were gathered from the literature, with preference given to those calculated at locations which were geographically proximate to our study areas [42,[49][50][51] (Fig 1E). We were unable to locate South African studies for hyena and used instead data from Kenyan populations [42]. ...
... Specific estimates for each predator-prey dyad were gathered from the literature, with preference given to those calculated at locations which were geographically proximate to our study areas [42,[49][50][51] (Fig 1E). We were unable to locate South African studies for hyena and used instead data from Kenyan populations [42]. Only for cheetah were we unable to find capture success values disaggregated by prey species and instead used overall success rate across all observed prey captures [50]. ...
Large mammalian herbivores use a diverse array of strategies to survive predator encounters including flight, grouping, vigilance, warning signals, and fitness indicators. While anti-predator strategies appear to be driven by specific predator traits, no prior studies have rigorously evaluated whether predator hunting characteristics predict reactive anti-predator responses. We experimentally investigated behavioral decisions made by free-ranging impala, wildebeest, and zebra during encounters with model predators with different functional traits. We hypothesized that the choice of response would be driven by a predator’s hunting style (i.e., ambush vs. coursing) while the intensity at which the behavior was performed would correlate with predator traits that contribute to the prey’s relative risk (i.e., each predator’s prey preference, prey-specific capture success, and local predator density). We found that the choice and intensity of anti-predator behaviors were both shaped by hunting style and relative risk factors. All prey species directed longer periods of vigilance towards predators with higher capture success. The decision to flee was the only behavior choice driven by predator characteristics (capture success and hunting style) while intensity of vigilance, frequency of alarm-calling, and flight latency were modulated based on predator hunting strategy and relative risk level. Impala regulated only the intensity of their behaviors, while zebra and wildebeest changed both type and intensity of response based on predator traits. Zebra and impala reacted to multiple components of predation threat, while wildebeest responded solely to capture success. Overall, our findings suggest that certain behaviors potentially facilitate survival under specific contexts and that prey responses may reflect the perceived level of predation risk, suggesting that adaptive functions to reactive anti-predator behaviors may reflect potential trade-offs to their use. The strong influence of prey species identity and social and environmental context suggest that these factors may interact with predator traits to determine the optimal response to immediate predation threat.
... Several studies stressed out the importance of vegetation structure and cover, which interact with the predator hunting technique and the prey escape technique to ultimately affect the hunting outcome. This was shown in savanna (Funston et al., 2001;Hopcraft et al., 2005) and temperate ecosystems (Husseman et al., 2003;Lone et al., 2014). Other habitat characteristics can interfere with the predator hunting success. ...
... However, to approach their prey undetected, ambush predators need concealment. Therefore, their hunting success is prone to be influenced by vegetation (Funston et al., 2001;Lone et al., 2014), and negatively affected by an increased aridity and a subsequent decrease in vegetation density (scenario 3, climatic conditions influence prey vulnerability through changes in habitat characteristics only). Whereas prey escape tactics might not be that crucial for the outcome of an encounter with an ambush predator, a climate-driven decrease in prey body condition will negatively influence prey fleeing capacity leading to an overall disadvantage against cursorial predators (scenario 2). ...
... When changes in climatic conditions influence prey vulnerability through changes in habitat characteristics and habitat-driven hunting success increases with rainfall (scenario 3), predator growth rates are negatively impacted by below average rainfall and positively influenced by above average rainfall. This is typically the case in African savannas where grass height is related to rainfall (Rutherford, 1980) and lions are hunters that are more successful in tall grass (Funston et al., 2001). Over the long term, predator populations benefit from the combined linear increase of prey abundance (Supplementary Figure S9) and non-linear increase of hunting success (i.e. ...
Climate changes affect the distribution and abundance of organisms, often via changes in species interactions. Most animals experience predation, and a number of models have investigated how climate fluctuations can influence predator–prey dynamics by affecting prey abundance through changes in resource availability. However, field studies have shown that prey vulnerability is a key feature determining the outcome of predator–prey interactions, which also varies with climatic conditions, via changes in prey body condition or in habitat characteristics (e.g. vegetation cover). In this theoretical work, we explore, with large mammals of African savannas in mind, how the interplay between climate-induced changes in prey abundance and climate-induced changes in prey vulnerability affects the immediate and long-term responses of predator populations. We account for prey body condition and habitat effects on prey vulnerability to predation. We show that predictions on how predator abundance responds to climate fluctuations differ depending on how climate influences prey vulnerability (habitat characteristics vs. prey body condition). We discuss how species traits influence the relative importance of the different sources of vulnerability. For example, our results suggest that populations of cursorial predators (such as spotted hyaenas) are expected to fare better than populations of ambush predators (such as African lions) in African ecosystems that will be characterised by an aridification. This study highlights the importance of understanding, and accounting for, the vulnerability factors associated to a given predator–prey pair, and improves our comprehension of predator–prey relationships in a changing climate.
... Therefore, under the niche divergence hypothesis, increased sexual dimorphism in carnivorous carnivorans is assumed to reduce intraspecific dietary competition by reducing competition between the sexes. Supporting this assumption are observations finding that females and males of many carnivoran species utilize different dietary resources [30][31][32][33] . ...
... Because male territories are larger and overlap with multiple smaller female territories 52-54 , male conspecifics exhibit larger range sizes and potentially encounter a wider range of prey. As a result, male conspecifics can exploit these novel, often larger prey items [30][31][32][33]55,56 due to morphological and functional adaptations (e.g. larger bite forces, stronger skulls and teeth) associated with their larger body sizes 13,14,[57][58][59] . ...
... Species of both clades are largely polygynous, and under sexual selection, larger males are more successful in defending territories against other males [60][61][62] . With larger territories, male conspecifics encounter and exploit different, often larger prey [30][31][32][33]63 by presumably using their larger bodies, skulls, teeth, and biting abilities to capture and consume these different prey 14,57-59 . Because all felids and musteline weasels are hypercarnivorous and specialize on vertebrate prey, niche divergence is hypothesized to reduce intraspecific competition for food by further maintaining size dimorphism in these traits 14,57,58 . ...
Although sexual size dimorphism (SSD) is widespread across the animal tree of life, the underlying evolutionary processes that influence this phenomenon remains elusive and difficult to tease apart. In this study, I examined how social system (as a proxy for sexual selection) and diet (as a proxy for natural selection) influenced the evolution of SSD in terrestrial carnivorans (Carnivora; Mammalia). Using phylogenetic comparative methods, I found that are territorial solitary and carnivorous carnivorans exhibited selection towards increased degree of male-biased SSD compared to other carnivorans with alternative social systems and diets. I also found the absence of Rensch’s rule across most carnivoran clades, suggestion a relaxation of the influences of sexual selection on SSD. These results together suggest that sexual selection and niche divergence together are important processes influencing the evolution of male-biased SSD in extant terrestrial carnivorans.
... We also hypothesized that the time of day could influence perceived predation risk, with most kills by lions occurring at twilight or night (Hayward & Slotow, 2009). Finally, we hypothesized that larger groups of lions would pose a greater risk to prey species than smaller groups, given hunting success is related to lion hunting group size (Funston et al., 2001;van Orsdol, 1984). We used the population of lions reintroduced to Addo Elephant National Park (Addo; Figure 1; Hayward, Adendorff, et al., 2007) to test these predictions. ...
... We cannot rule out a lion-focused scenario whereby lions approach their favored prey downwind thereby negating potential avoidance by prey ( Figure 2). There is mixed evidence of this as several studies conclude lions exhibit no clear preference for hunting or increased hunting success in a particular wind direction (Funston et al., 2001;Stander & Albon, 1993;van Orsdol, 1984), whereas Wijers et al. (2022) found that lions hunted in crosswind conditions. Furthermore, it seems highly unlikely that lions would choose when to mate and fight, for example, based on their distance from prey, or would choose to situate themselves further from prey species they preferentially hunt and kill, than nonpreferred prey. ...
Predators can induce behavioral changes in prey that influence vigilance, grouping patterns, and space use, and these can ultimately affect prey demography and trophic interactions. Consequently, prey must respond to the risk of predation, but little is known about the features that drive the spatial responses of prey species to predators. We tested what factors affected the proximity of prey to the lions reintroduced to Addo Elephant National Park, South Africa. We also tested whether prey species that are preferentially killed by lions revealed greater responsiveness than those that are not, and whether prey respond to predator behavioral states and hunger. From 1588 observations of potential prey locations in relation to lions under varying wind directions , lion behaviors, and hunger states throughout the day and night, we found no evidence of wind-driven odor responses affecting prey proximity to lions. Prey species that were not preferentially preyed upon by lions occurred closer to lions than those species that lions prefer to hunt. Prey were closer to lions performing noisy behaviors compared to those performing quiet behaviors. Prey were more likely to be closer to covertly behaving lions and further from stationary lions. Our results, compared to the published literature and accepted dogma of the primacy of odor in predator detection, suggest large vertebrate prey responses to predators in intact, multi-species assemblages are context dependent. K E Y W O R D S avoidance behavior, olfaction, predation risk, prey preferences, vigilance
... Only waterbuck had an affinity for areas with low visibility, corresponding with their aforementioned preference for dense riverine vegetation also seen in other studies (Owen-Smith, 1996;Smit et al., 2007). Greater grass height and vegetation cover are important for predator hunting success (Funston et al., 2001;Owen-Smith, 2019), so many species may use open areas as an anti-predator strategy (Funston et al., 2001;Hopcraft et al., 2005). Buffalo, eland and zebra used more open areas, but are not preferred prey species of lions in Majete (Briers-Louw & Leslie, 2020), possibly due to their success in avoiding lion predation by using the areas of higher visibility. ...
... Only waterbuck had an affinity for areas with low visibility, corresponding with their aforementioned preference for dense riverine vegetation also seen in other studies (Owen-Smith, 1996;Smit et al., 2007). Greater grass height and vegetation cover are important for predator hunting success (Funston et al., 2001;Owen-Smith, 2019), so many species may use open areas as an anti-predator strategy (Funston et al., 2001;Hopcraft et al., 2005). Buffalo, eland and zebra used more open areas, but are not preferred prey species of lions in Majete (Briers-Louw & Leslie, 2020), possibly due to their success in avoiding lion predation by using the areas of higher visibility. ...
Understanding ecological-and management-related predictors of mammal space use within protected areas is critical for management planning. This is particularly true in small, fenced and isolated reserves where we hypothesised that management-related activities will influence large herbivore space use more than ecological characteristics. We used camera trap data to assess the space use patterns for 18 ungu-late species in the small and isolated Majete Wildlife Reserve (Majete) in the understudied Miombo woodland ecoregion of southern Malawi. In the 2018 dry season, the 691 km 2 reserve was systematically surveyed for ungulate presence at 140 camera trap locations. Over a period of 5456 camera days, the survey yielded 11 078 independent detections of 18 ungulate species and three predators. Using a single-species occupancy modelling framework, the probability of space use of ungulates was assessed in relation to five management (fire exposure, fire frequency , water availability, distance from fence and road) and five ecological (visi-bility, grass biomass, vegetation type, terrain ruggedness and predator abundance) space use covariates, while accounting for imperfect detection. Top-ranked models contained multiple covariates for 15 of the 16 species modelled, with only nyala's (Tragelaphus angasii) space use best predicted by vegetation type only. Distance to water, vegetation type, visibility and fire frequency were predictors having strong influences on six or more species each. More management-related covariates reflected in top models, but ecological covariates had more meaningful effect sizes making us reject the hypothesis. Importantly though, distance from the roads and fence were also identified as prominent predictors. Notably, black rhinoceros' (Diceros bicornis) probability of space use increased with distance from the reserve boundary. Beyond informing habitat management for ungulates in Majete, these results can form the basis for understanding species-specific space use patterns in other small reserves with similar characteristics and threats.
... Thus, lions have the advantage in closed habitats while hyenas are likely to benefit from open habitats due to their cursorial nature. Lions tend to select for habitats with tall grass or steep embankments that promote hunting success and increases the catchability of prey [58][59][60], while hyenas seemingly appear to be able to utilize any type of habitat as habitat generalists [38,39]. ...
... As mainly cursorial predators, spotted hyenas are generally more active, travel faster and further than lions [38,53]. As sit-and-wait predators, lions typically have lower activity, move at slower speeds, and cover shorter distances than hyenas [40,59]. Corresponding to the findings of Durant et al. [77], lions and hyenas in this study utilized similar habitats, both mainly occurring within grassland habitats in Etosha and shrubland habitats in Botswana [106][107][108]. ...
Resource partitioning promotes coexistence among guild members, and carnivores reduce interference competition through behavioral mechanisms that promote spatio-temporal separation. We analyzed sympatric lion and spotted hyena movements and activity patterns to ascertain the mechanisms facilitating their coexistence within semi-arid and wetland ecosystems. We identified recurrent high-use (revisitation) and extended stay (duration) areas within home ranges, as well as correlated movement-derived measures of inter- and intraspecific interactions with environmental variables. Spatial overlaps among lions and hyenas expanded during the wet season, and occurred at edges of home ranges, around water-points, along pathways between patches of high-use areas. Lions shared more of their home ranges with spotted hyenas in arid ecosystems, but shared more of their ranges with conspecifics in mesic environments. Despite shared space use, we found evidence for subtle temporal differences in the nocturnal movement and activity patterns between the two predators, suggesting a fine localized-scale avoidance strategy. Revisitation frequency and duration within home ranges were influenced by interspecific interactions, after land cover categories and diel cycles. Intraspecific interactions were also important for lions and, important for hyenas were moon illumination and ungulates attracted to former anthrax carcass sites in Etosha, with distance to water in Chobe/Linyanti. Recursion and duration according to locales of competitor probabilities were similar among female lions and both sexes of hyenas, but different for male lions. Our results suggest that lions and spotted hyenas mediate the potential for interference competition through subtle differences in temporal activity, fine-scale habitat use differentiation, and localized reactive-avoidance behaviors. These findings enhance our understanding of the potential effects of interspecific interactions among large carnivore space-use patterns within an apex predator system and show adaptability across heterogeneous and homogeneous environments. Future conservation plans should emphasize the importance of inter- and intraspecific competition within large carnivore communities, particularly moderating such effects within increasingly fragmented landscapes.
... Likewise, we analyzed whether standardized bill length is related to hummingbird hunting strategy by calculating Pearson's contingency coefficient. 57 The analyses were performed in the InfoStat program. 52 ...
... There are variables that allow us to evaluate their consumption, such as prey availability, morphological features, habitat structure (prey detectability), territorial behavior, social hierarchy, and feeding preferences. 57,58 Hummingbirds depend on flower patches for food, which is related to the phenology of flowering plants. Although, hummingbirds have been considered primarily nectarivores, it has been reported that some species may include insects at their diet, some only use this resource in certain seasons and others only consume insects. ...
An organism's morphological features can determine the type of prey they consume due to adaptive advantages in capturing them, for example, the shape and length of the bill in birds. Hummingbirds have been considered specialists in nectar consumption. However, they have been documented to also be important insectivores in ecosystems. In this study, we evaluated arthropod consumption by hummingbirds in relation to their bill and capture strategies. We theorized that hummingbirds with short-medium bills (9-11 bill length to body mass cube root ratio) capture a higher proportion of non-flying arthropods, as they have a lower linear speed of closure of the bill tip relative to the base, while species with long bills (13-14) capture mainly flying arthropods. The study was conducted in El Triunfo Biosphere Reserve, Chiapas, southeastern Mexico from 2015 to 2016, where seven hummingbird species that captured arthropods, their strategies of capture, and the type of prey consumed were recorded. We also analyzed the stomach contents (n = 72 stomachs) collected in the Sierra Madre de Chiapas. We calculated the frequency of occurrence of arthropods and their diversity, as well as the breadth of the feeding niche and the bill-prey relationship of 15 hummingbird species. The Green-throated Mountain-gem ( Lampornis viridipallens) and Rivoli's Hummingbird ( Eugenes fulgens) presented the highest richness of arthropod consumption. Arthropod prey belonged to eight orders, where Hymenoptera and Diptera were the most frequent. The most recorded capture strategies were on the substrate in sustained flight (37.5%) and hovering in sustained flight (33.3%). A positive association was found between the type of arthropods consumed and the morphological traits of the hummingbirds, non-flying arthropods were captured by hummingbirds with short-medium bills, while hummingbirds with long bills captured flying arthropods. Morphological adaptations for food resource use are a trait that determines food selection, capture, and handling success.
... We previously used playback experiments in Amboseli National Park, Kenya to demonstrate that individuals within a social group can benefit directly from the influence of an older leader because of their enhanced ability to make crucial decisions about predators [4]. While elephant family groups within a stable, natural study population were consistently able to assess the greater risk associated with three roaring lions versus one, those with older matriarchs were significantly better at the more subtle task of identifying the increased threat associated with male versus female roars (male lions being more than 50% larger and at a distinct advantage in capturing large-bodied prey [36,38,39]). However, ecological knowledge may be affected by experience, as well as age, particularly given that many free-ranging elephant populations have been subjected to severe human disturbance through habitat loss, culling and poaching [22]. ...
... Bigger groups of hunting lions are invariably more successful in taking large-bodied prey such as elephants [18,19,35,36,38,39]. Discriminating between the numbers of roaring lions is-under normal circumstances-likely to be a universal skill among elephant family groups that is acquired at a relatively early stage of adulthood [4]. ...
The transmission of reliable information between individuals is crucial for group-living animals. This is particularly the case for cognitively advanced mammals with overlapping generations that acquire detailed social and ecological knowledge over long lifetimes. Here, we directly compare the ecological knowledge of elephants from two populations, with radically different developmental histories, to test whether profound social disruption affects their ability to assess predatory threat. Matriarchs (≤50 years of age) and their family groups received playbacks of three lions versus a single lion roaring. The family groups in the natural Amboseli population (Kenya) reliably assessed the greater predatory threat presented by three lions roaring versus one. However, in the socially disrupted Pilanesberg population (South Africa), no fine-scale distinctions were made between the numbers of roaring lions. Our results suggest that the removal of older and more experienced individuals in highly social species, such as elephants, is likely to impact the acquisition of ecological knowledge by younger group members, particularly through the lack of opportunity for social learning and cultural transmission of knowledge. This is likely to be exacerbated by the trauma experienced by juvenile elephants that witnessed the culling of family members and were translocated to new reserves. With increasing levels of anthropogenic disturbance, it is important that conservation practitioners consider the crucial role that population structure and knowledge transfer plays in the functioning and resilience of highly social and long-lived species.
... Empirical insight into group foraging has centred upon model vertebrate systems such as mammals (Creel & Creel, 1995;Fanshawe & Fitzgibbon, 1993;Funston et al., 2001;Holekamp et al., 1997), birds (Beauchamp, 2012;Cook et al., 2017;Harel et al., 2017;Krebs et al., 1972) and fish (Grand & Dill, 1999;Harpaz & Schneidman, 2020;Johnsson, 2003;Pitcher et al., 1982), or invertebrates at the extremes of social organisation (i.e. eusocial ants and bees; Balbuena et al., 2012;Bockoven et al., 2015). ...
... These effects may be amplified in environments where the immediate foraging space scales weakly (or not at all) with the size of foraging groups, as is the case in our experimental arenas and the log interiors inhabited by velvet worms (Barclay et al., 2000a). Such density effects are broadly aligned with previous findings in E. rowelli (Reinhard & Rowell, 2005), as well as studies of archetypal social foragers such as African wild dogs (Creel & Creel, 1995;Fanshawe & Fitzgibbon, 1993), lions (Funston et al., 2001) and spotted hyenas (Holekamp et al., 1997). ...
Group living is ubiquitous in nature, and social foraging is among the common forms of cooperative behaviour in animals. Understanding the evolution and maintenance of social foraging requires knowledge of the underlying benefits and costs to group members, though these are only known for few model taxa, which often lie at the extremes of social organisation. Here, we experimentally examined hypothesised benefits and costs of social foraging in the velvet worm Euperipatoides rowelli; the only Onychophoran currently known to live and forage in groups. To explore benefits, we tested the effects of natural group size and controlled prey size on the latency to attack and begin consuming prey, the likelihood of complete prey consumption and the time taken to completely consume prey. Our focal cost was aggressive intraspecific interactions in the form of biting, kicking and striking, which may present a particular risk to soft-bodied velvet worms. As predicted, we found a positive scaling of most effects with the size of foraging groups. Larger groups were faster to attack and begin consuming prey, and more likely to completely consume prey, though they took longer to do so. This may be a consequence of the increasing representation of juveniles in larger aggregations. However, larger groups were also subject to heightened aggression among conspecifics, though is it unclear whether such a cost holds at the level of individual group members. Variation in prey size and, hence, resource availability had little effect across all outcomes, except for a slightly reduced likelihood of completely consuming larger prey. A time-course analysis of individual activity suggested the existence of feeding hierarchies with adults excluding juveniles, particularly when resources are scarce. Taken together, our results offer the first support for the existence of foraging benefits and costs to group membership in E. rowelli, which may contribute to the maintenance of group living in this evolutionary significant taxon.
... First, the lunar phase (full moon) during nocturnal observations, for although lions do not alter their movement activities under moonlight (Preston et al., 2019), prey species often do, with those relying on non-visual cues typically reducing activity (Prugh & Golden, 2014), and others increasing their use of high lion-encounter areas (wildebeest and zebra) or congregating in greater abundance in low-risk areas (buffalo and gazelle; Palmer et al., 2017). African lions also have higher hunting success on nights when the moon is absent or obscured, in open (Funston et al., 2001;Packer et al., 2011;Van Orsdol, 1984) and wooded areas (Preston et al., 2019). Therefore, although the proportion of all observed hunts (n = 52, including by unknown lions) that were successful here (25%) is similar to previous observations (Elliot et al., 1977;Van Orsdol, 1984), inclusion of data from dark nights might have increased observed success (Funston et al., 2001). ...
... African lions also have higher hunting success on nights when the moon is absent or obscured, in open (Funston et al., 2001;Packer et al., 2011;Van Orsdol, 1984) and wooded areas (Preston et al., 2019). Therefore, although the proportion of all observed hunts (n = 52, including by unknown lions) that were successful here (25%) is similar to previous observations (Elliot et al., 1977;Van Orsdol, 1984), inclusion of data from dark nights might have increased observed success (Funston et al., 2001). Whether dark night hunting success is due to higher encounter rate, attack rate or attack success is unclear, but this study's low bright-moon nocturnal encounter probability might partially underlie this pattern, with important consequences (Packer et al., 1990). ...
Predation is a fundamental ecological process influencing the distribution and abundance of animal populations and underlying how prey species perceive risk. The predation process is composed of four sequential stages – search, encounter, attack and kill – each of which has been used to describe risk across the landscape. Here, we used direct observational data of free‐ranging, radio‐collared African lions in Serengeti National Park's western corridor to (1) investigate daily and seasonal predation stage probabilities and (2) using two analytical approaches, compare four mechanisms – prey distribution, intra‐specific competition, spatially anchored landscape features and predator hunting method – that potentially drive spatial predation stage patterns. Results showed that lions encountered potential prey at night significantly less than during diurnal or crepuscular periods. Nocturnal observations were predominantly during full‐moon phases, so if this lower nocturnal encounter rate was due to moon phase it may contribute to lions' typically poor full‐moon hunting success. Predation stage probabilities did not differ between seasons despite high variability in seasonal prey abundance. Spatially, lions encountered potential prey in prey‐rich, open areas near water and spatial range centres. Compared with available areas within seasonal ranges, lion attacks were more likely where prey abundance was high, and kill locations were associated with prey‐rich areas near water and range centres, collectively suggesting opportunistic hunting. However, compared with preceding predation stage locations, attacks occurred near range peripheries and kills where hunting cover was greater, suggesting ambush predation. Our results indicate substantial temporal and spatial variation across the different stages of the predation process. They also highlight first, that results can vary in important ways depending on how analyses are approached, and second, that understanding predator‐prey dynamics depends on analyses of the different stages of predation. This study used direct observational data of African lions in Serengeti National Park's western corridor to (1) investigate daily and seasonal predation stage probabilities and (2) compare, using two analytical approaches, four mechanisms ‐ prey distribution, intra‐specific competition, spatially anchored landscape features and predator hunting method ‐ that potentially drive spatial predation stage patterns. Results indicate substantial temporal and spatial variation across the different stages of the predation process, and highlight that results can vary in important ways depending on how analyses are approached, and that understanding predator‐prey dynamics depends on analyses of the different stages of predation.
... Given that lions are more likely to hunt successfully if they are able to approach prey within 20 m (Orsdol 1984), we defined a group as the number of cattle within a 20 m radius of the focal individual and recorded the number of cattle within the focal individual's group at the conclusion of each 10-min observation period. Because vegetation height has been found to correlate with lion hunting success (Funston et al. 2001), we also estimated the average height of vegetation at each observation point. Finally, given the tendency of lion attacks to increase during evening hours (Orsdol 1984), i.e. after sunset (approximately 06:30 p.m. during this study), we recorded the time of day at the conclusion of each 10-min observation period (Altmann et al. 1973, Djagoun et al. 2013. ...
... We also found that cattle in both high and low risk village rangelands formed smaller groups as vegetation height increased. This may seem counter-intuitive as well, as vegetation height and density is typically associated with lion hunting success (Orsdol 1984, Funston et al. 2001, Davies et al. 2016). However, we observed this pattern regardless of level of depredation risk. ...
Fear of predators fundamentally shapes the ecology of prey species and drives both inter‐ and intra‐specific interactions. Extensive research has examined the consequences of predation risk from large carnivores on the behavior of wild ungulate prey species. However, many large carnivores not only hunt wild prey but also depredate domestic livestock, especially in pastoralist systems where livestock share land and resources with large carnivores. Northern Tanzania is a hotspot for human‐carnivore conflict driven by livestock depredation and interactions are particularly severe between African lions (Panthera leo) and pastoralist cattle (Bos taurus). In this ecosystem, we explored the degree to which pastoralist cattle exhibited anti‐predator behaviors during their daily grazing routines. Using focal animal sampling, we compared two typical anti‐predator behaviors, vigilance and grouping, among cattle in village rangelands with high and low background depredation rates. We found that cattle in high risk village rangelands formed 21.2% larger groups than cattle in low risk village rangelands. Interestingly, cattle in low risk village rangelands spent 68.4% more time vigilant than cattle in high risk village rangelands. These patterns were influenced significantly by the time of day: as sunset approached, cattle in low risk village rangelands spent more time vigilant and cattle in high risk village rangelands formed larger groups. These results suggest that pastoralist cattle exhibit anti‐predator strategies that vary both spatially and temporally, and that such strategies might help livestock optimally trade‐off the costs and benefits of anti‐predator behavior across timescales (i.e., the risk allocation hypothesis). We discuss the implications of our results for husbandry techniques that might reduce behavioral costs associated with cattle anti‐predator behaviors and help increase tolerance for lions and other large carnivores. These improvements are critical to human‐carnivore coexistence given the prevalence of pastoralism globally and the rising potential for conflict with large carnivores such as lions.
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... Specifically, species that rely on visual acuity for cursorial hunting techniques (sensu African wild dog Lycaon pictus; Rasmussen and Macdonald, 2011) are more active during moonlit nights. Conversely, the success of species that rely on ambush strategies (sensu African lion, Panthera leo) is either unaffected (Cozzi et al., 2012) or reduced (Funston et al., 2001;Packer et al., 2011;Preston et al., 2019) by the presence of moonlight, and these species are therefore often less active on bright nights. Variation in predator response to moonlight has a knock-on effect on prey species, resulting in cyclical variation in the level of predation risk associated with nocturnal foraging across the lunar cycle (Owen-Smith, 2019;Tyler et al., 2016;Wu et al., 2018). ...
... Furthermore, lion are the single significant predator of giraffe, both generally (Berry, 1973;Strauss and Packer, 2013) and in our study site (Stander et al., 2018), and the hunting efficiency of lion is reduced on moonlit nights (e.g. Funston et al., 2001). Considering their high visual acuity (Veilleux and Kirk, 2014), increased light intensity at night likely improves both predator detection and foraging efficiency for giraffe (Prugh and Golden, 2014). ...
Natural cycles of light and darkness shift the balance of risks and gains for animals across space and time. Entrainment to photic cycles allows animals to spatiotemporally adapt their behavioural and physiological processes in line with interplaying ecological factors, such as temperature, foraging efficiency and predation risk. Until recently, our understanding of these chronobiological processes was limited by the difficulties of 24 hr observations. Technological advances in GPS biotelemetry however are now allowing us unprecedented access to long-term, fine-scale activity data. Here we use data derived from frontline technology to present the first large-scale investigation into the effects of natural fluctuations of light and darkness on the locomotor activity patterns of a threatened African mega-herbivore, the giraffe (Giraffa spp.). Using data from a remote population of Angolan giraffe (G. g. angolensis) in the northern Namib Desert, Namibia, we reveal the first full picture of giraffe chronobiology in a landscape of fear. Furthermore, we present clear evidence of the effect of moonlight on the nocturnal activity patterns of large ungulates. Our results are in line with recent research demonstrating that, rather than a fixed internal representation of time (circadian clock), many surface-dwelling ungulates have plastic activity patterns that are vulnerable to modification by external factors including light and temperature. Relatedly, we highlight important conservation management implications of rising temperatures and increasing light pollution on the chronobiology of surface-dwelling mammals.
... Excluding cheetah encounters, for which we had a limited sample size to discuss inferences, encounters peaked for all dyads during periods of high moonlight illumination. Given that not all guild species' activity levels are influenced by moonlight availability (Cozzi et al., 2012), this may reflect the lower risks associated with approaching potentially contested resources during periods of high illumination, for example reduced lion ambush risks because of greater visibility (Funston et al., 2001). Alternatively, encounter peaks across moonlight levels may have been driven by periodicity in the use of shared areas of home ranges, which in turn, may have been driven by periodicity in resource distributions (Riotte-Lambert, Benhamou & Chamaill e-Jammes, 2013). ...
... Interestingly, leopard-lion and leopard-wild dog encounters also showed secondary peaks during periods of low light availability. This may simply reflect the impacts of low light levels on species detectability (Funston et al., 2001). In other words, encounters may have increased during these periods because species were able to travel closer to other guild members, with or without intent, without being detected and species remained unaware of one another's presence. ...
Encounters between individuals can have implications for a range of processes, including disease transmission, information transfer and competition. For large carnivores, difficulties in directly observing individuals and historical hardware limitations of GPS collars mean that relatively little is known of the spatio‐temporal factors contributing to encounters. The African large predator guild represents one of the few remaining functionally intact guilds of large carnivores on the globe and so represents a unique study system for understanding competitor interactions. We explored the drivers of male leopard ( Panthera pardus ) encounters with lions ( Panthera leo ), African wild dogs ( Lycaon pictus ) and cheetahs ( Acinonyx jubatus ) in the context of habitat characteristics and temporal activity overlaps. Using high‐resolution (five minute GPS fixes) data from 48 large African carnivores from 2012 to 2018, we quantified encounter occurrences between male leopards and other guild species and related these to habitat type (open vs closed), activity overlaps and moonlight levels. Leopards met wild dogs 4.56 ± 1.15 (standard error), lions 3.11 ± 0.56 and cheetahs 2.27 ± 0.73 times per month. All species instigated encounters, but leopard instigated encounters with dominant competitors appeared to reflect imperfect information on risk, primarily occurring within habitats with limited visibility. Moreover, encounters peaked during periods of high temporal overlap, suggesting that, although previous research indicates temporal activity patterns may not be driven by predator avoidance, temporal overlap has implications for competitor dynamics. Our results show how habitat characteristics and niche overlaps contribute to encounters between competitors and provide an example of how niche shifts within competitor assemblages can impact competition between species.
... As each hour passes in the model, animals move to nearby locations weighing vegetation preferences and a fear of lions (based on a recent lion kill in a location). At the same time, lions move towards nearby animals and if a lion encounters an animal, then the lion has a probability of making a kill (Funston et al., 2001) at that location. The amount of time each herbivore agent spends on the various vegetation types and the number of lion kills in each location are recorded by the model and output after 30 days have passed in the simulation. ...
One of the primary goals of the palaeosciences is to produce robust understandings of palaeoecologies of extinct ecosystems. The time has arrived where such palaeoecologies can be significantly improved—agent-based models (ABMs) that synthesize our modern understandings of animal ecology with past conditions provide a unique opportunity for this. The Palaeo-Agulhas Plain (PAP), a now submerged landscape off the south coast of South Africa, formed a novel ecosystem during the lower sea levels of the Pleistocene. Here we provide a review of the utility of ABMs for understanding herbivores in prehistoric environments using the Paleoscape Model. The Paleoscape Model is fortunate to have features models of climate, geology and vegetation upon which we can understand how herbivores used the PAP environment. To incorporate aspects of herbivore behaviour and habitat suitability into the model requires analogies from extant systems. In addition to reconstructions of the availability of resources and associated risks, data on herbivore behaviour and physiology are imperative to our understanding of these systems. Behaviour, driven by changes in the environment, is interpreted by species, leading to selection of suitable habitats ultimately driving the herbivore’s decisions within ABMs. Finally, we discuss herbivore integration into these models that can be used in numerous other scenarios (past, present and future).
... As each hour passes in the model, animals move to nearby locations weighing vegetation preferences and a fear of lions (based on a recent lion kill in a location). At the same time, lions move towards nearby animals and if a lion encounters an animal, then the lion has a probability of making a kill (Funston et al., 2001) at that location. The amount of time each herbivore agent spends on the various vegetation types and the number of lion kills in each location are recorded by the model and output after 30 days have passed in the simulation. ...
One of the primary goals of the palaeosciences is to produce robust understandings of palaeoecologies of extinct ecosystems. The time has arrived where such palaeoecologies can be significantly improved-agent-based models (ABMs) that synthesize our modern understandings of animal ecology with past conditions provide a unique opportunity for this. The Palaeo-Agulhas Plain (PAP), a now submerged landscape off the south coast of South Africa, formed a novel ecosystem during the lower sea levels of the Pleistocene. Here we provide a review of the utility of ABMs for understanding herbivores in prehistoric environments using the Paleoscape Model. The Paleoscape Model is fortunate to have features models of climate, geology and vegetation upon which we can understand how herbivores used the PAP environment. To incorporate aspects of herbivore behaviour and habitat suitability into the model requires analogies from extant systems. In addition to reconstructions of the availability of resources and associated risks, data on herbivore behaviour and physiology are imperative to our understanding of these systems. Behaviour, driven by changes in the environment, is interpreted by species, leading to selection of suitable habitats ultimately driving the herbivore's decisions within ABMs. Finally, we discuss herbivore integration into these models that can be used in numerous other scenarios (past, present and future).
... The goal of cooperation is to increase the success rate in problemsolving and to perform activities that are difficult to complete by a single individual. For instance, in many species, the hunting success rate is significantly higher when animals cooperate (Hector 1986;Funston et al. 2001), while others, through cooperation, are able to hunt prey that is difficult to catch (Ford et al. 1998;MacNulty et al. 2014). Cooperative hunting is probably the most common type of cooperation (Packer and Ruttan 1988), and it has been observed even in non-social animals (Dinets 2014;Lührs and Dammhahn 2010), and between different species (Eaton 1969;Bayley and Rose 2020). ...
Dolphins live in a fission-fusion society, where strong social bonds and alliances can last for decades. However, the mechanism that allows dolphins to form such strong social bonds is still unclear. Here, we hypothesized the existence of a positive feedback mechanism in which social affiliation promotes dolphins' cooperation, which in turn promotes their social affiliation. To test it, we stimulated the cooperation of the 11 dolphins studied by providing a cooperative enrichment tool based on a rope-pulling task to access a resource. Then we measured the social affiliation [simple ratio index (SRI)] of each possible pair of dolphins and evaluated whether it increased after cooperation. We also evaluated whether, before cooperation, pairs that cooperated had a higher SRI than those that did not cooperate. Our findings showed that the 11 cooperating pairs had significantly stronger social affiliation before cooperation than the 15 non-cooperating pairs. Furthermore, cooperating pairs significantly increased their social affiliation after cooperation, while non-cooperating pairs did not. As a result, our findings provide support to our hypothesis, and suggest that the previous social affiliation between dolphins facilitates cooperation, which in turn promotes their social affiliation.
... Animals that typically avoid human disturbance may be more likely to take risks, such as foraging on anthropogenic food sources, when their body condition is poor and their options are limited. For example, drought-induced reduction in vegetation cover makes it harder for ambush predators like lions (Panthera leo) to hunt 40 . Combined with reduced prey availability, this change in habitat structure and prey accessibility led to an increase in lion attacks on livestock in Botswana during drought 41 . ...
Climate change and human–wildlife conflict are both pressing challenges for biodiversity conservation and human well-being in the Anthropocene. Climate change is a critical yet underappreciated amplifier of human–wildlife conflict, as it exacerbates resource scarcity, alters human and animal behaviours and distributions, and increases human–wildlife encounters. We synthesize evidence of climate-driven conflicts occurring among ten taxonomic orders, on six continents and in all five oceans. Such conflicts disrupt both subsistence livelihoods and industrial economies and may accelerate the rate at which human–wildlife conflict drives wildlife declines. We introduce a framework describing distinct environmental, ecological and sociopolitical pathways through which climate variability and change percolate via complex social–ecological systems to influence patterns and outcomes of human–wildlife interactions. Identifying these pathways allows for developing mitigation strategies and proactive policies to limit the impacts of human–wildlife conflict on biodiversity conservation and human well-being in a changing climate. The authors summarize the underappreciated role of climate change in amplifying human–wildlife conflict. They synthesize evidence of climate-related conflicts and introduce a framework highlighting the environmental, ecological and sociopolitical pathways linking climate change to conflict outcomes.
... In the wild, ecological factors such as activity of prey and predation risk potentially lead to different activity and hunting peaks between lions and cheetahs [93][94][95]. In zoos, obviously, hunting became obsolete for cheetahs and lions. ...
Mammals are constantly exposed to exogenous and endogenous influences that affect their behaviour and daily activity. Light and temperature, as well as anthropogenic factors such as husbandry routines, visitors, and feeding schedules are potential influences on animals in zoological gardens. In order to investigate the effects of some of these factors on animal behaviour, observational studies based on the analyses of activity budgets can be used. In this study, the daily and nightly activity budgets of six lions (Panthera leo) and five cheetahs (Acinonyx jubatus) from four EAZA institutions were investigated. Focused on the influencing factor light and feeding, we analysed these activity budgets descriptively. Behaviour was recorded and analysed during the winter months over an observation period of 14 days and 14 nights using infrared-sensitive cameras. Our results show that lions and cheetahs exhibit activity peaks at crepuscular and feeding times, regardless of husbandry. Thus, lions in captivity shift nocturnal behaviour familiar from the wild to crepuscular and diurnal times. In cheetahs, in contrast, captive and wild individuals show similar 24hr behavioural rhythms. The resting behaviour of both species is more pronounced at night, with cheetahs having a shorter overall sleep duration than lions. This study describes the results of the examined animals and is not predictive. Nevertheless, the results of this study make an important contribution to gaining knowledge about possible factors influencing the behaviour of lions and cheetahs in zoos and offer implications that could be useful for improving husbandry and management.
... Indeed, lion translocations sometimes occur across multiple countries or regions and are becoming increasingly common throughout Africa (Wolf and Ripple, 2018;Bertola et al., 2022). There are a variety of motivations for translocations, including restoration of locally extirpated populations or augmentation of existing populations, human-lion conflict mitigation, reestablishing populations for tourism purposes, and for a range of other commercial, personal, ethical, and financially motivated reasons (Table 1; Stander, 1990;Funston et al., 2001;Hunter et al., 2007;Trinkel et al., 2008;Miller and Funston, 2014;Morapedi et al., 2021). Substantial guidance exists to ensure translocations benefit the conservation of species and habitats (IUCN/SSC, 1998;Peŕez et al., 2012;IUCN/SSC, 2013;IUCN/ SSC, 2018;Soorae, 2018;Berger-Tal et al., 2019). ...
As the top predator in African ecosystems, lions have lost more than 90% of their historical range, and few countries possess strong evidence for stable populations. Translocations (broadly defined here as the capture and movement of lions for various management purposes) have become an increasingly popular action for this species, but the wide array of lion translocation rationales and subsequent conservation challenges stemming from poorly conceived or unsuitable translocations warrants additional standardized evaluation and guidance. At their best, translocations fill a key role in comprehensive strategies aimed at addressing the threats facing lions and fostering the recovery of wild populations in their historic range. At their worst, translocations can distract from addressing the major threats to wild populations and habitats, divert scarce funding from more valuable conservation actions, exacerbate conflict with humans in recipient sites, disrupt local lion demography, and undermine the genetic integrity of wild lion populations in both source and recipient sites. In the interest of developing best practice guidelines for deciding when and how to conduct lion translocations, we discuss factors to consider when determining whether a translocation is of conservation value, introduce a value assessment for translocations, and provide a decision matrix to assist practitioners in improving the positive and reducing the negative outcomes of lion translocation.
... The mean distance goats travelled from the kraal was less than the mean distance of all predation events from the kraals, suggesting that they did indeed occupy a relatively safe zone with respect to predators. Similarly, goats were generally released at least two hours after cattle and returned at least two hours before cattle in the afternoon, hence temporally avoiding natural peak lion hunting periods that occur from dusk till dawn (Funston, Mills and Biggs, 2001). Perhaps this sense of security renders goats at greater risk of predation on the rare occasions that predators do venture close to kraals. ...
Human-carnivore conflicts occur globally and are a leading cause of carnivore population declines. Such conflicts usually occur when carnivores predate livestock and can include preemptive and retaliatory killing of carnivores by livestock farmers. In northern Botswana, livestock farming is a widespread and culturally important practice. Subsistence farming enterprises commonly abut protected areas, and human-carnivore conflicts are common. Understanding interactions between livestock and carnivores, and how livestock use resources and habitats generally, are important components to managing these conflicts. Throughout this thesis, I explore human-carnivore conflict in northern Botswana. I found that livestock resource selection and predation vary seasonally and spatially in relation to ecological and anthropogenic features in the landscape. Predation sites are subsequently avoided by cattle in the short-term, but not by goats. Contemporary mitigation to minimise livestock predation events commonly includes lethal control and broadscale exclusion by artificial barriers and aversive interventions, yet naturally occurring deterrent signals fine-tuned through evolution are rarely considered. Lions roar to deter conspecifics from territorial boundaries, which prey and subordinate carnivores eavesdrop on and modify their movement and behaviour in response. I used lion vocalisations to understand livestock (prey) responses to this apex carnivore and to test how effective roars are in deterring lions and other carnivores. Using a high-tech experimental approach, I found that (1) cattle avoid lion vocalizations, while goats do not, and (2) lions are not deterred by lion roars played-back from Remotely Operated Acoustic Repellent stations (ROARs), nor are other human-carnivore conflict species occurring in the area. Finally, I used a commonly occurring anti-predator signal in nature, demonstrating that artificial eyespots painted on cattle rumps deter lions from attacking cattle. Collectively, the results from my thesis can be used to better manage livestock in a landscape of risk, and to promote human-carnivore coexistence by deterring predation. Applications derived from my thesis to promote human-carnivore coexistence can be used across Africa and the globe.
... Ungulates should aggregate in areas with high-quality and abundant forage, but foraging theorists suggest ungulate herding behavior is also an adaptation to predation risk, where gregariousness improves predator detection and decreases individual time spent on vigilance (Lima and Dill 1990;Kie 1999). Both mixed feeders and browsers are also expected to encounter greater predation risk in areas with dense woody vegetation or tall grass where predator detection is inhibited, compared to grazers in more open habitats (Funston et al. 2001;Fritz and Loison 2006;Valeix et al. 2009). ...
Habitat selection is a dynamic biological process where species respond to spatiotemporal variation in resource availability. The resulting distribution patterns can be detected as presence–absence or heterogeneity in abundance and indicate habitat preferences based on environmental correlations at multiple scales. Variation in habitat selection by ungulates is constrained by trade-offs in top-down and bottom-up trophic processes arising from differences in forage requirements, water dependency, anthropogenic effects, and predation avoidance, and mediated by physiological (feeding guild) and morphological (body size) factors. We conducted distance sampling over 7 years in the Tarangire Ecosystem (TE) of northern Tanzania for six resident ungulate species: Kirk’s dik-dik (Madoqua kirkii), Grant’s gazelle (Nanger granti), Thomson’s gazelle (Eudorcas thomsonii), Masai giraffe (Giraffa camelopardalis tippelskirchi), impala (Aepyceros melampus), and common waterbuck (Kobus ellipsiprymnus), and tested hypotheses related to effects of top-down and bottom-up processes on ungulate presence and abundance. We modeled ecological correlates against two distributional responses to understand which environmental factors constrained these ungulate species at different scales; (i) presence–absence observations modeled in a logistic regression to assess habitat selection at an ecosystem scale; (ii) local abundances from presence-only observations modeled using a negative binomial distribution for finer-scale selection. Browser and grazer species in the TE selected suitable habitat proximal to rivers and avoided the Combretum–Azanza woody plant assemblage. Browsers and grazers also showed strong preference for habitat with more dense cover of preferred forage species, and abundance was influenced by the presence of specific forage species with significant seasonal variation. Mixed feeders were more heterogeneous in habitat suitability implying that broader diets allow avoidance of areas with high human activity. Small-bodied and dehydration-sensitive species selected areas near rivers and seasonal tributaries. Seasonal habitat selection was more pronounced among mixed feeders. Conservation strategies based on spatially and seasonally explicit resource selection studies such as ours can minimize impacts to biodiversity by protecting vital resources to ungulates through all seasons of the year.
... First, movement can be difficult in the wet season due to extensive flooded regions and the prevalence of thick grasses up to 3 m in height which would increase energetic costs. Second, movement through tall grass impedes the ability of wild dogs to scan for risks (and prey) and increases the risk of predation from stalking predators such as lions [25]. Thus, it is possible that decreasing movement during the wet season is a mechanism to conserve energy and reduce predation risk. ...
Background
Prey depletion is a threat to the world’s large carnivores, and is likely to affect subordinate competitors within the large carnivore guild disproportionately. African lions limit African wild dog populations through interference competition and intraguild predation. When lion density is reduced as a result of prey depletion, wild dogs are not competitively released, and their population density remains low. Research examining distributions has demonstrated spatial avoidance of lions by wild dogs, but the effects of lions on patterns of movement have not been tested. Movement is one of the most energetically costly activities for many species and is particularly costly for cursorial hunters like wild dogs. Therefore, testing how top-down, bottom-up, and anthropogenic variables affect movement patterns can provide insight into mechanisms that limit wild dogs (and other subordinate competitors) in resource-depleted ecosystems.
Methods
We measured movement rates using the motion variance from dynamic Brownian Bridge Movement Models (dBBMMs) fit to data from GPS-collared wild dogs, then used a generalized linear model to test for effects on movement of predation risk from lions, predictors of prey density, and anthropogenic and seasonal variables.
Results
Wild dogs proactively reduced movement in areas with high lion density, but reactively increased movement when lions were immediately nearby. Predictors of prey density had consistently weaker effects on movement than lions did, but movements were reduced in the wet season and when dependent offspring were present.
Conclusion
Wild dogs alter their patterns of movement in response to lions in ways that are likely to have important energetic consequences. Our results support the recent suggestion that competitive limitation of wild dogs by lions remains strong in ecosystems where lion and wild dog densities are both low as a result of anthropogenic prey depletion. Our results reinforce an emerging pattern that movements often show contrasting responses to long-term and short-term variation in predation risk.
... In addition, their tree-climbing abilities (with or without a kill; Balme et al. 2017a) also allow leopards to coexist with spotted hyaenas and lions in the same area, without the need for spatiotemporal partitioning. However, spotted hyaenas require more expansive, open areas to run down prey (Kruuk 1972;Périquet et al. 2014), whereas lions require vegetation cover to stalk their prey (Funston et al. 2001;Hopcraft et al. 2005). Even though habitat type did not significantly influence spatial partitioning on Madikwe, it was an important influence on occupancy in our second-best model, and thus it is still an important influence to consider when evaluating large carnivore space use. ...
Context: The spatio-temporal partitioning of large carnivores is influenced by interspecific competition and coexistence within small, enclosed reserves. Lions (Panthera leo), spotted hyaenas (Crocuta crocuta) and leopards (Panthera pardus) are the three largest African carnivores and have the greatest potential for intra-guild competition, particularly where space is limited.
Aim: To investigate the spatio-temporal partitioning between lions, spotted hyaenas and leopards in a small (~75 000 ha), enclosed nature reserve, Madikwe Game Reserve (Madikwe), South Africa.
Methods: We deployed 110 camera traps (baited n = 55 and unbaited n = 55) across Madikwe from 26 August 2019 until 6 May 2020. Von Mises kernel density plots were used to investigate daily temporal partitioning among the three species. A multiple-species, single-season occupancy model was used to investigate daily space use patterns.
Key results: We found both temporal and spatial exclusion between lions and spotted hyaenas on Madikwe. However, no evidence was found of spatio-temporal partitioning between lions and leopards, and spotted hyaenas and leopards.
Conclusions: Exploitative and interference competition on Madikwe might be high enough to warrant spatio-temporal partitioning between lions and spotted hyaenas to avoid the negative effects of intra-guild competition. Contrastingly, patterns observed between leopards and both lions and spotted hyaenas preclude the possibility of top-down control by superior carnivores.
Implication: These findings call for an adaptive management approach, where both carnivore and prey species compositions are constantly monitored. Management strategies such as these will allow for the conservation of valuable resources (i.e. prey species) to ensure the persistence of large carnivore populations across African ecosystems.
... Zebras Equus quagga foraged in areas burnt Biological Reviews (2022) annually or triennially during the day, but avoided triennially burnt areas at night, as they contained more dense woody vegetation favouring ambush predators (e.g. lions), which are more active at night (Funston, Mills & Biggs, 2001). Similarly, impala Aepyceros melampus foraged in triennially burnt and unburnt areas during the day, but used less risky areas at night (i.e. annually burnt areas) when the risk of lion predation was greater (Burkepile et al., 2013). ...
Both fire and predators have strong influences on the population dynamics and behaviour of animals, and the effects of predators may either be strengthened or weakened by fire. However, knowledge of how fire drives or mediates predator-prey interactions is fragmented and has not been synthesised. Here, we review and synthesise knowledge of how fire influences predator and prey behaviour and interactions. We develop a conceptual model based on predator-prey theory and empirical examples to address four key questions: (i) how and why do predators respond to fire; (ii) how and why does prey vulnerability change post-fire; (iii) what mechanisms do prey use to reduce predation risk post-fire; and (iv) what are the outcomes of predator-fire interactions for prey populations? We then discuss these findings in the context of wildlife conservation and ecosystem management before outlining priorities for future research. Fire-induced changes in vegetation structure, resource availability, and animal behaviour influence predator-prey encounter rates, the amount of time prey are vulnerable during an encounter, and the conditional probability of prey death given an encounter. How a predator responds to fire depends on fire characteristics (e.g. season, severity), their hunting behaviour (ambush or pursuit predator), movement behaviour, territoriality, and intra-guild dynamics. Prey species that rely on habitat structure for avoiding predation often experience increased predation rates and lower survival in recently burnt areas. By contrast, some prey species benefit from the opening up of habitat after fire because it makes it easier to detect predators and to modify their behaviour appropriately. Reduced prey body condition after fire can increase predation risk either through impaired ability to escape predators, or increased need to forage in risky areas due to being energetically stressed. To reduce risk of predation in the post-fire environment, prey may change their habitat use, increase sheltering behaviour, change their movement behaviour, or use camouflage through cryptic colouring and background matching. Field experiments and population viability modelling show instances where fire either amplifies or does not amplify the impacts of predators on prey populations, and vice versa. In some instances, intense and sustained post-fire predation may lead to local extinctions of prey populations. Human disruption of fire regimes is impacting faunal communities, with consequences for predator and prey behaviour and population dynamics. Key areas for future research include: capturing data continuously before, during and after fires; teasing out the relative importance of changes in visibility and shelter availability in different contexts; documenting changes in acoustic and olfactory cues for both predators and prey; addressing taxonomic and geographic biases in the literature; and predicting and testing how changes in fire-regime characteristics reshape predator-prey interactions. Understanding and managing the consequences for predator-prey communities will be critical for effective ecosystem management and species conservation in this era of global change.
... Lions hunt mostly at night (Fischhoff et al. 2007;Hayward and Hayward 2007;Hayward and Slotow 2009) and less often during the day (Van Orsdol 1984). Lions rely upon the cover provided by tall grass, dense woody vegetation, or gullies to ambush their prey (Funston et al. 2001;Hopcraft et al. 2005;Loarie et al. 2013;Davies et al. 2016). They may also concentrate their hunting around water sources that draw aggregations of ungulates (Davidson et al. 2012(Davidson et al. , 2013. ...
Movement patterns of lions (Panthera leo) reveal how they hunt large herbivores in heterogeneous landscapes such as the Kruger National Park in South Africa. Large herbivores are distributed differently on the landscape and therefore have different vulnerabilities as prey for lions. For instance, blue wildebeest (Connochaetes taurinus) occupy small grazing lawns at night but are difficult for lions to capture because open areas lack cover for stalking. African buffalo (Syncerus caffer) aggregate in large herds but are less available because these herds only intermittently enter the home ranges of individual lion prides. Unlike large herds of wildebeest and buffalo, plains zebra (Equus quagga) move widely in small herds while browsing greater kudus (Tragelaphus strepsiceros) and giraffes (Giraffa camelopardalis giraffa) generally occur in lower densities. We used spatial data derived from GPS collars to investigate several hypotheses regarding the movements of three lion prides in response to their prey. We found that lions were most active and moved longer distances during nighttime than during daytime. Lions remained within their core home ranges on 87% of nights and wandered to the outlying areas of the home ranges every second night. Lions visited grazing lawns, that is, area of short grass, where wildebeest herds resided every second night, and moved toward the direction of buffalo herds within 2 km of vicinity. Lions spent more time near riverbanks that provided dense woody cover at night than expected but concentrated only weakly near sites with surface water where herbivores drank in the dry season. Our study contributes to understanding how lions vary their movements in response to the spatial and temporal heterogeneity in the relative availability and vulnerability of multiple prey species.
... Part of this competition most likely originates from the stress experienced by juvenile and subadult males when pride males or other adult lions harassed them. Having abundant food is likely to reduce these incidences, alleviating influences that typically impact subadult survival including reduced hunting success (Funston, Mills & Biggs, 2001) and opportunities to acquire territories (Funston et al., 2003) when emigrating from natal areas (Hanby & Bygott, 1987). The observations that there were relatively more abundant subadult males 18 months post-drought suggest that juvenile and subadult males survived better during the drought. ...
... Predation threat can also vary with changes in night-time illumination (Harmsen et al. 2011;Penteriani et al. 2011Penteriani et al. , 2013Palmer et al. 2017). We found that dingo activity around waterholes increased on moonless nights, suggesting that dingo hunting success may improve under low light conditions, as has been shown for other savanna predators (e.g., lions: Funston et al. 2001;Packer et al. 2011). This is because prey may find detection or defence against dingoes more difficult on moonless nights, particularly when young are present. ...
When exotic species are introduced to new environments, they often have a competitive advantage over native species. In northern Australia, pigs, cattle, and water buffalo have established widespread, feral populations. As ungulates have high water requirements, they typically congregate near waterpoints. We used a fencing experiment to test whether native macropods preferentially visited savanna waterholes where large ungulates were excluded. We also investigated whether water scarcity affected the visitation behaviour and temporal activity patterns of herbivores at waterholes and whether increasing prey aggregation at waterholes increased dingo presence. We found that macropods did not use fenced waterholes preferentially over unfenced ones. Cattle presence at waterholes increased as water became scarce, while macropod and pig presence peaked in the middle of the dry season. Macropod activity declined rapidly at the end of the dry season when cattle activity was greatest, suggesting that macropods may avoid waterholes in areas utilised by cattle when competition for resources is high. Macropods and all ungulates visited waterholes more during a drought year compared to an average rainfall year. Despite increasing prey activity, dingo presence at waterholes did not increase when water became scarce. However, dingo presence increased significantly on moonless nights. Our results suggest that competition between macropods and ungulates may intensify during periods of water scarcity. Climate change and pastoral intensification are likely to increase competition for resources between ungulates and macropods in Australian savannas, potentially threatening macropod populations across the landscape in the future.
Significance statement
In northern Australia, feral populations of pigs, cattle, and water buffalo compete with native wildlife for access to water sources. As interspecific competition favours species with a size advantage, we tested whether kangaroos and wallabies (macropods) preferentially use waterholes where large ungulates (cattle and buffalo) were excluded. We found that macropods avoided waterholes when cattle presence was high but did not preferentially use waterholes where livestock were excluded. When water scarcity peaked during a drought, macropods and all three feral ungulate species visited waterholes more. However, increased prey presence at waterholes during the drought did not correspond with increased predator (dingo) presence. Our study advances the understanding of behavioural interactions between invasive and native species at important shared resources, and how this may affect wildlife conservation in an increasingly unpredictable environment.
... avec des effectifs estimés entre 1800 et 4000 individus, les populations de lions d'afrique centrale et occidentale sont les plus menacées, mais ont été jusqu'ici très peu étudiées. par ailleurs, alors qu'en afrique orientale et australe l'organisation sociale du lion africain a fait l'objet de nombreuses recherches (schaller, 1972 ;stander, 1991 ;Funston et al., 2001 ;packer et al., 2005), peu d'études ont été menées sur ce sujet en afrique occidentale et centrale. Or Bauer et al. (2003) émettent l'hypothèse que le schéma d'organisation sociale tel que décrit classiquement pourrait ne pas s'appliquer entièrement aux populations d'afrique centrale et de l'ouest, en parallèle avec une différence de régime alimentaire observée entre ces régions. ...
L’«Étude Lion Zakouma» est née au début de l’année 2003 dans le but de participer à mieux connaître les populations de lions (Panthera leo) d’Afrique centrale, jusqu’ici peu étudiées, alors qu’elles semblent de plus en plus menacées par la diminution des habitats naturels et l’augmentation parallèle des conflits Homme-prédateurs. La zone d’étude choisie correspond à la moitié est du Parc national de Zakouma (PNZ) – réserve naturelle située dans le Sud-est du Tchad – et couvre une surface d’environ 1.500 km². Elle englobe la majorité des points d’eau permanents du parc et abrite la plus grande part de la faune présente dans le PNZ en saison sèche. Les activités de terrain de l’«Étude Lion Zakouma» se sont déroulées pendant la saison sèche, d’avril 2003 à juin 2006, et se sont concentrées sur les objectifs suivants : l’identification de la population de lions, l’analyse de son régime alimentaire, l’évaluation de son statut sanitaire, l’étude des conflits Homme-prédateurs en périphérie du parc et la formation d’agents locaux aux tâches de suivi de la population de lions. La taille de la population de lions du PN Z a été estimée à l’aide des techniques de «reconnaissance individuelle» et de recensement nocturne par «call-in», à environ 140 individus (dont 63 ont été identifiés individuellement). Elle peut être considérée comme une petite population mais elle fait vraisemblablement partie d’une métapopulation au sein de laquelle des échanges génétiques sont possibles. Cinq lions ont été équipés de colliers émetteurs VHF puis suivis par radio-tracking aérien et terrestre afin de rassembler des informations sur la structure sociale, la reproduction et la prédation de cette population. L’aire de prédilection de ces cinq lions pendant la saison sèche a également été déterminée ainsi que son évolution au fil des campagnes de terrain. Le taux de reproduction des femelles et les caractéristiques démographiques de la population correspondent a ce qui a été observé ailleurs. Le taux de survie des lionceaux semble par ailleurs très satisfaisant. Trois coalitions de mâles résidants et un mâle résidant apparemment solitaire, couvrant le territoire d’au moins cinq groupes familiaux, ont été identifiés dans la zone d’étude entre 2003 et 2006. Le mode d’organisation des lions dans le PN Z semble néanmoins différer légèrement du schéma classique tel que décrit par Schaller (1972). Le «suivi continu» de la prédation est unanimement considéré comme la technique de choix pour déterminer avec précision le régime alimentaire des lions. Du fait de la densité du couvert végétal limitant visibilité et déplacements en véhicule dans notre zone d’étude, il ne nous a pas été permis de l’utiliser. À sa place, nous avons choisi la technique du «suivi occasionnel», qui donne un bon aperçu des habitudes alimentaires des carnivores mais tend à entraîner une surreprésentation des proies de grande taille. L’Éléphant (Loxodonta africana), le Buffle (Syncerus caffer brachyceros), le Bubale (Alcephalus buselaphus lelwel) et le Cobe defassa (Kobus ellipsyprimnus defassa) font partie des espèces dont la prédation a été la plus observée dans la zone d’étude. Les trois premières sont également les espèces les plus représentées dans le parc. Des prélèvements de sang ont été réalisés sur les lions anesthésiés pour la pose d’un collier émetteur et ont fait l’objet d’analyses biologiques. Les virus associés à la maladie du carré, l’immunodéficience féline (FIV), la panleucopénie féline (typhus du chat), la calicivirose, la péritonite infectieuse féline (PIF), la leucose féline (FeLV ) et la rhinotrachéite ont été recherchés. Ces analyses n’ont révélé aucune menace sur la santé des lions du PN Z. Divers moyens ont été utilisés pour évaluer l’importance des conflits Homme-prédateurs dans la zone périphérique du PNZ. Parmi ceux-ci, une enquête portant sur la prédation touchant le bétail domestique et les moyens mis en place pour la prévenir
a été menée auprès d’éleveurs transhumants et sédentaires appartenant à onze villages et six campements
nomades de la zone périphérique du parc. Si la prédation sur le bétail s’est avérée être un problème régulier
dans quelques villages et de nombreux campements nomades, l’ensemble des éleveurs consultés semblent considérer jusqu’à maintenant les dégâts liés à la prédation comme bénins par rapport aux pertes liées à la maladie et au vol. Les différentes activités de l’ «Étude Lion Zakouma» ont été menées en synergie avec le «Volet Suivi écologique» du projet CURESS (projet européen en charge de la gestion du PNZ à l’époque de notre étude) et ses différents agents ont été formés à travers une instruction théorique et pratique aux différentes techniques de recensement et de suivi des lions du parc.
... In addition, organisms can forage socially (e.g., as part of a cohesive group), can either forage for themselves or 'scrounge' food from other group members (Giraldeau and Dubois, 2008), and can adopt different foraging modes (e.g., actively search vs 'sit and wait') (Visser and Fiksen, 2013). Lions, for example, may sometimes hunt food solitarily, but usually forage in a group known as a 'pride' (Funston et al., 2001). Interestingly, a group of foraging crows is called a 'murder' (Bergan, 2015). ...
Foraging by animals is such an important and fundamental aspect of animal behaviour, that we need to understand it as well as possible, which means we need to be successful in our attempts to explain and predict observed foraging behaviour. The best way to achieve such success is to invoke evolutionary theory with animals hypothesised to forage in ways that maximise their biological fitness.
This evolutionary approach leads to Optimal Foraging Theory (OFT), which includes Classic OFT, if individual animals do not respond directly to the foraging behaviour of others, and the more-general Foraging Game Theory, if such responses do occur. In either case, the basic hypothesis is that animals in a population will adopt a foraging strategy such that no individual can achieve higher biological fitness by deviating from the rest of the population. If individuals do not respond to the foraging behaviour of others, this hypothesis is equivalent to classic maximisation of some currency that acts as proxy for biological fitness. If such responses do occur, then individuals are collectively involved in foraging ‘games’ that are expected to be at equilibrium.
The foraging process involves different stages (e.g., pre- and post-ingestion) and a number of types of decision (e.g., patch choice, diet choice, patch exploitation, movements, group membership, producer vs scrounger, foraging mode), leading to studies that focus on such different foraging stages and decisions. Studies of foraging behaviour have consequently focused on one or other of these stages and decisions.
OFT has grown and developed enormously during its 50-year history, through expansion, extension, application and inspiration, with observations generally supporting expectations, to the point where it is now a ‘strong theory’ of behaviour and ecology. Understanding foraging behaviour remains an area of much interest and research, as befits its fundamentally important role in all of life.
The articles in the section on Foraging in this Encyclopedia illustrate these general aspects of foraging in that they rely on evolutionary theory to understand observed foraging behaviour, embrace OFT in particular, describe how OFT has grown and developed, and indicate current research interest. They also focus on the foraging stages and decisions outlined above.
... Further, many animals that form socially stable groups also have sub-groups of individuals that hunt for the group, which has been observed in several well-studied mammal groups. For instance, male lions, and the females of some chimpanzee (Pan trogolodytes) groups rarely participate in hunts (Boesch, 1994;Funston, Mills, & Biggs, 2001;Funston, Mills, Biggs, & Richardson, 1998;Scheel & Packer, 1991) -though membership is still highly predictable in these hunting groups. ...
... This area is adjacent to open savannah and woodland habitat directly next to the Great Ruaha River, which provides the only yearround natural source of water for wildlife in the park used by giraffes and other prey (Mtahiko et al., 2006). We suspect that lions may be using hunting grounds near water to increase hunting success (sensu Funston et al., 2001;Spong, 2002). However, lion hunting behaviour and giraffe availability do not alone explain why giraffes are highly selected prey for lions in Ruaha National Park. ...
Rates at which predators encounter, hunt and kill prey are influenced by, among other things, the intrinsic condition of prey. Diseases can considerably compromise body condition, potentially weakening ability of afflicted prey to avoid predation. Understanding predator–prey dynamics is particularly important when both species are threatened, as is the case with lions (Panthera leo) and giraffes (Giraffa camelopardalis). Importantly, an emergent disease called giraffe skin disease (GSD) may affect predatory interactions of lions and giraffes. Hypotheses suggest that GSD may negatively affect the likelihood of giraffes surviving lion attacks. We evaluated giraffe–lion interactions in Ruaha National Park, Tanzania, where 85% of the giraffe population has GSD. We monitored lion hunting behaviour and estimated proportion of the giraffe population with GSD and evidence of ‘lion marks’ from assumed previous lion predation attempts (i.e. claw marks, bite marks and missing tails). Although we recorded lions hunting and feeding on 16 different prey species, giraffes represented the largest prey category (27%; n = 171 of 641). For age and sex cohorts combined, 26% (n = 140 of 548) of encountered giraffes displayed evidence of previous lion predation attempts. Occurrence of lion marks was higher for adults and males in the giraffe population, suggesting that these individuals were more likely to survive lion attacks. We also found marginal evidence of a positive relationship between giraffes with severe GSD and occurrence of lion marks. Our results identify giraffes as important prey species for lions in Ruaha National Park and suggest that GSD severity plays a minor role in likelihood of surviving a lion attack. This is the first study to explore connections between lion predation and GSD. We explore the ecological implications of disease ecology on predator–prey interactions and consider opportunities for future research on causal links between GSD and giraffe vulnerability to lion predation. Diseases can considerably compromise body condition and movement of prey species, potentially weakening the ability of afflicted individuals to avoid predation. We evaluated giraffe–lion interactions in Ruaha National Park, Tanzania, where 85% of the giraffe population has giraffe skin disease (GSD), and estimated the proportion of the giraffe population with GSD and evidence of previous lion predation attempts. Our results identify giraffes as important prey for lions in Ruaha National Park and suggest that GSD plays a minor role in likelihood of giraffes surviving a lion attack.
... Lions can survive on a broad range of prey species that vary between habitats (Hayward and Kerley 2005, Borrego et al. 2018, van Hooft et al. 2018. Its prey consists mainly of mammals particularly ungulates weighing 190-550 kg with a preference for wildebeest (Connochaetes taurinus), plains zebra (Equus burchelli), and African buffalo (Syncerus caffer) (Mills and Shenk 1992, Funston et al. 2001, Kissui and Packer 2004, and warthog (Scheel andPacker 1991, Hayward andKerley 2005). Although lions are most active at night, they frequently hunt during the day (Schaller 1972). ...
The demography of the African lion is increasingly shaped by interactions with humans. Habitat fragmentation and persecution by humans are both linked to the decline in lions in most of their historical ranges such that current populations are largely restricted to isolated protected areas. This study examined the spatial and temporal patterns of lion killings in the Maasai steppe ecosystem. We used eighty-two lion mortality records for the last 13 years (2005-2017). Distances from the roads, river, lake, settlements, and the Normalized difference vegetation index value extracted for each lion killing location were the key landscape variables used to map the lion anthropogenic mortalities. There was a significant difference (p<0.05) between female and male lions killed from 127 mortality records. The anthropogenic retaliatory killing caused 77.9% of female and 22.1% of male mortalities. About 58% of the lions killed were adults, 39.1% were sub-adults and only 2.9% were cubs. The majority of lion killings incidences took place during the wet season around the Maasai homestead. The lion killings incidences were rampant in the eastern side but slightly clustered in the northern part. Vegetation cover in the actual lion killings areas influenced lion killings incidences. Distances from the public roads, rivers, and human settlements significantly (p<0.05) contributed to lion anthropogenic mortalities. It is anticipated that retaliatory killings of lions could intensify due to growing cattle herds in the ecosystem. To promote coexistence between humans and lions, conservation authorities should invest more in awareness and sensitization programs on the conservation of lions.
... Lions also exhibit sexually selected infanticide where newcomer males kill unrelated infants (Bertram, 1975;Packer and Pusey, 1983b;Chakrabarti and Jhala, 2019), thus resident males aggressively defend against invaders not only to maintain access to females but also to prevent the loss of progeny and fitness (Grinnell et al., 1995). Males occasionally capture prey large enough to feed the entire pride, but females perform the bulk of hunting, which mostly involves small-to medium-sized prey (Schaller, 1972;Funston et al., 2001), and, hence, males frequently scavenge and parasitize from kills made by the pride females (Schaller, 1972). Thus, within lion prides, males and females perform separate tasks-males: territorial defense and protection from infanticidal males, females: food, territorial defense, and rearing of cubs. ...
Understanding sexual segregation is crucial for comprehending sociality. A comparative analysis of long-term lion data from Serengeti and Ngorongoro in Tanzania, and Gir in India, reveals that male-female associations are contingent upon male and female group size, prey- size and availability, and the number of prides that each male coalition currently resides. Males maintain proximity with females, while females are responsible for segregation except at large kills. Lions feed on the largest prey in Ngorongoro and smallest in Gir, and females spend the most time with males in Ngorongoro and the least in Gir. Females roar less often in prey-scarce circumstances in Serengeti and throughout the year in Gir possibly to prevent being tracked by males that parasitize on female kills. However, females readily associate with males when available prey is large and abundant. Contrasting availability of resources between Gir and Serengeti/Ngorongoro helps explain varying degrees of sexual segregation and appears to drive differences in mating systems between these lion populations.
... The film and script Despite well-researched and long-standing estimates of hunting ability and success levels (e.g. Funston et al., 1998Funston et al., , 2001, the failure of many of the pride's hunts is presented as an unusual and looming threat to pride survival rather than something that is routine to all prides. Other existential drama includes one of the young males confronted by a large pack of hyenas. ...
Natural history documentaries are a globally important source of information about wildlife, conservation and environmental issues, and they are the closest many will get to seeing featured animals and their behaviour in the wild. They are entertainment, certainly, but may also inform people's knowledge of the natural world and influence their ideas on conservation of species and habitats. We locate our perspective in the existing literature analysing wildlife documentary making and its effects.
We argue that a conspicuous pre‐occupation with the ‘personalisation’ of individual animals and the injection of false jeopardy in recent wildlife documentaries leads to significant misinformation and creates problems for public understanding of wider conservation.
We illustrate our point by detailing episodes from the BBC natural history series Dynasties, discussing personalisation, anthropomorphism and the use of jeopardy to gain emotive impact and audience engagement. We find that narratives are framed around a single individual, that ‘stories’ are framed as soap operas, that jeopardy is emphasised throughout and that animals are endowed with the capacity to be aware of, and work towards, the dynasties of the title.
With conservation increasingly relying on public support, we argue that it is important that people are presented with factually correct information, and portraying wild animals as soap opera style characters is neither honest nor helpful.
A free Plain Language Summary can be found within the Supporting Information of this article.
... On the other hand, temporal processes, such as timing of daily photoperiods or the cyclicity of environmental temperatures, can impact animals' ability to obtain available resources through their effect on functional performance (e.g., visual sensory ability; Bosiger and McCormick 2014). For example, terrestrial carnivores often demonstrate relatively high activity in peripheral habitats, where preys are more abundant (Šálek et al. 2010;Loarie et al. 2013), but also tend to occupy these habitats at night, when hunting success is highest due to decreases in the preys' ability to detect and evade predators (Funston et al. 2001;Hopcraft et al. 2005;Loarie et al. 2013). Similar dependencies of animal activity patterns on resource supply and animals' capability of using resources are documented across many terrestrial and aquatic taxa (Ciechanowski et al. 2007;Blair 2009;Carter et al. 2015). ...
How animals partition activity throughout the day is influenced by processes that affect supply and obtainability of resources. However, as resource supply and usability are often entrained by the same diurnal pattern, it has been difficult to disentangle their relative importance. Given the strong influence that tide has on the distribution and accessibility of resources, intertidal systems present opportunities to examine questions surrounding the drivers of activity patterns. Here, we used multisensory biologgers to study the activity patterns of a coastal marine predator, sicklefin lemon sharks (Negaprion acutidens), in a tidally driven environment. Hidden Markov models were used to identify relatively high and low locomotory activity states, which were used as proxies for behavioural–activity states and to examine the factors underpinning variation in activity patterns. Although tide governs the spatial distributions of this species and showed some effect on sharks’ activity, diurnal light patterns were the predominant factor influencing behavioural-activity patterns, with the probability of high activity peaking overnight. Temperature and body size also had minor negative influences on the probability of animals being in the high-activity state. Interestingly, sharks were least likely to be in a high-activity state during high tide, a time of presumed high resource supply, contradicting the common assumption that this species forages during high tide. We suggest that despite the importance of the accessibility of resources, functional constraints, such as sensory (e.g., visual) and mechanical (e.g., swimming) performance ultimately underpin the activity patterns of intertidal marine predators through their influence on foraging success.
... This is a reasonable assumption in some cases. For example, males and females do not differ in hunting success in wild dogs (Creel and Creel 1995) or lions (Funston et al. 2001). ...
Models of optimal group size need to identify the currency that correctly captures the fitness consequences of foraging. Although daily intake or daily net energy gain per animal are widely used as currencies, they are not ideal. They predict that all available time should be spent hunting and do not reflect performance during a hunt. We argue that the net rate while hunting is a better currency. Using an example based on the African wild dog, we illustrate the difference between maximizing daily net energy and net rate. Using the same example, we show that if foraging is limited by constraints on energy expenditure, then the optimal group size can be different from the size that maximizes the net rate while hunting. The direction of the effect depends on whether costs increase or decrease with group size. Furthermore, the proportion of time spent resting can be predicted. We suggest two novel approaches for future models: to consider the optimal hunting group size given a fixed group size and to investigate how the presence of dependent young may affect foraging behavior. We hope this will lead to meaningful conclusions on the role cooperative hunting has played in the evolution of sociality in social carnivores.
... Although this does not translate directly into hunting success, it does suggest that lions actively decide which prey to hunt . Furthermore, hunting skills associated with small pride size might also influence prey selection in the reserve (Bissett, Bernard & Parker, 2012;Packer et al., 1990;Funston, 2001). ...
Apex carnivores play an important role in the ecosystem by regulating prey via predation. Anthropogenic influences have resulted in rapid range and population reductions of large carnivores across the African continent. These carnivores are often reintroduced into protected areas to compensate for human-induced losses, restore ecosystem functioning and promote eco-tourism.
Majete Wildlife Reserve in Malawi is a prime example, as human persecution resulted in the extirpation of large carnivores, with the exception of a small spotted hyena (Crocuta crocuta; hereafter hyena) population. As from 2003, African Parks attempted to rectify this problem by restoring and developing the reserve. Between 2011 and 2012, three lions (Panthera leo) and six leopards (Panthera pardus) were reintroduced. The aim of this study was to describe the ecology of the apex predators and to determine whether the felid reintroduction was successful or not.
Lion and leopard movements and home ranges were determined using GPS collars. The reintroduction of felids was considered successful. This was based on: (1) reduced post-release movements; (2) lack of homing tendencies; (3) breeding success; and (4) population persistence. Mean home ranges of lion (380.45 ± 117.70 km2 [SE]) and leopard (495.08 ± 80.99 km2), were the largest on record for any reintroduced felid in Africa, which was likely due to a low competitor density. Thus, we expect home range sizes to decrease with an increase in conspecific density.
Population abundances and densities were estimated with the use of camera traps. The known lion population increased to eleven individuals in five years, while the leopard population was estimated at 11 (range = 9–17). This indicates population persistence and growth. Both founder populations were small and require additional translocations to maintain genetic diversity. Hyena density (2.62 hyenas/100 km2) and clan size (5.33 ± 0.67) were the lowest estimates in any woodland habitat and comparable to arid areas. This may be explained by decades of direct persecution and poaching of their prey, or a naturally low density.
Predator diets were described and compared by means of scat analysis. Lion and hyena exhibited a high dietary overlap of medium to large herbivores. Using Jacobs’ preference index, both species preferred warthog (Phacocoerus africanus) and waterbuck (Kobus ellipsiprymnus). Hyenas selected a broader range of prey, likely reducing competition with lions (which almost exclusively selected only four species). In contrast, leopards occupied a lower dietary niche, which consisted mainly of small-to medium-sized ungulates. These findings indicate that the three apex predators use resource partitioning to reduce competition.
This study suggests that reintroduction is a viable tool for re-populating large carnivores in protected areas in Malawi. The current predator population appeared to have a minimal impact on prey populations due to their small population size. We recommend long-term monitoring of predator-prey dynamics as the predator populations increase to prevent major ecological imbalances. Finally, we encourage management to focus energy and resources on the formation of a managed carnivore metapopulation to establish a genetically viable carnivore population within Malawi.
... However, foraging in areas rich in preferred prey does not necessarily translate into hunting success and thus learned response (i.e. adult lion(s) acquiring the skill to effectively hunt warthogs and waterbuck and teaching the technique to other pride members) associated with small pride size might also explain this prey selection by lions (Bissett et al., 2012;Funston, Mills, & Biggs, 2001;Packer, Schell, & Pusey, 1990). ...
Between 2011 and 2012, the carnivore guild in Majete Wildlife Reserve (MWR), Malawi, was restored following the reintroduction of lion ( Panthera leo ) and leopard ( Panthera pardus ). The aim of this study was to describe and compare the diet of lion, leopard and resident spotted hyaena ( Crocuta crocuta ) using scat analysis. Lions and spotted hyaenas displayed the greatest dietary overlap ( O ab = 0.88) and selected mainly medium‐ to large‐bodied prey species. Lions had a mean preferred prey weight of 120.33 ± 42.14 kg (SE), with warthog ( Phacochoerus africanus ) and waterbuck ( Kobus ellipsiprymnus ) making up 60.64% of relative biomass consumed. Spotted hyaenas had a mean preferred prey weight of 102.40 ± 41.69 kg and had a more generalised diet ( B a = 0.46) compared to lions ( B a = 0.36). In contrast, leopards occupied a dietary niche substantially lower than that of lions and spotted hyaenas, selecting relatively smaller prey with a mean preferred prey weight of 27.50 ± 6.74 kg. Our results suggest that coexistence between the resident hyaena and reintroduced lion and leopard in MWR is facilitated by dietary partitioning. We advise long‐term monitoring of reintroduced carnivores in small, enclosed reserves to assess their impacts on predator and prey populations.
... Temporal partitioning across the diel cycle may be partially explained in the context of hunting (Cozzi et al., 2012). For ambush predators, such as lions and leopards, low-light conditions increase hunting success (Funston, Mills, & Biggs, 2001;Packer, Swanson, Ikanda, & Kushnir, 2011;Martins & Harris, 2013). Therefore, nocturnal activity levels may remain consistent across moon phases due to the need to meet minimum energetic requirements and the limited hunting opportunities available across other diel periods. ...
Understanding the mechanisms facilitating coexistence within species assemblages is a key consideration for conservation as intact assemblages are necessary for maintaining full ecosystem function. The African large predator guild represents one of the few remaining functionally intact large predator assemblages on Earth, and as such, represents a unique study system to understand competitive interactions. Yet, relatively little is known of the coexistence mechanisms between some of its intermediately sized members, particularly leopards ( Panthera pardus ). Here, we use overlapping spatio‐temporal activity and GPS data on lions ( Panthera leo ), leopards, African wild dogs ( Lycaon pictus ) and cheetahs ( Acinonyx jubatus ) to examine spatial interactions and temporal partitioning between leopards and other guild members in northern Botswana. We found that at the population level, male leopard space use and activity patterns were largely unaffected by intraguild competitors. Leopards showed minimal movement coherence with competitors (avoidance or attraction) when moving through areas of home ranges shared with intraguild species. Moreover, we found evidence to support the hypothesis that guild species’ activity patterns are primarily driven by light availability rather than predator avoidance. Our results suggest predator avoidance has a limited impact on broad‐scale leopard spatio‐temporal niches, with aspects of the leopards’ ecology and life history likely facilitating its ability to thrive in close proximity to competitors. Considered alongside other studies, our results suggest that landscape‐level approaches to conservation may be suitable for aiding leopard conservation.
... National Park in Namibia, Stander, 1991). Lion social dynamics in undisturbed populations predicts lower survival rate of sub-adult males resulting from various risks encountered when emigrating from natal areas (Hanby & Bygott, 1987) such as reduced hunting success (Funston, Mills, & Biggs, 2001) or reduced opportunities to acquire territories (Funston et al., 2003). Apparent survival rates for adult females were significantly higher than that of adult males. ...
Changes in demographic features of the lion population in the south‐western Kgalagadi Transfrontier Park cause conservation concern. The ratio between adult males and females changed from 1♂:2♀ to 1♂:1♀ over a 32‐year period (1977–2010). This is atypical for undisturbed lion populations. We evaluated mechanisms that on their own or together could explain the trend by using both a discrete‐ and continuous‐time capture–recapture model to analyse 2 years of individual‐based observation data (May 2013–June 2015). Although most vital rates were within expected parameters, shifts in the sex structure remained similar to that reported for sub‐adults previously. The population comprised of 26.9% cubs and juveniles (<2 years), 14.2% sub‐adults (2–4 years) and 59% adults (>4 years). Litters had equal sex ratios at first detection. Birth rates were lower (0.44 cubs per female per year) than between 1998 and 2001 (0.69 cubs per female per year). The continuous model indicated a higher survival effect for female cubs, while both models showed higher survival rates/effects for sub‐adult and adult females relative to males. However, the sex ratio for sub‐adults was male‐biased. The shift in demographic signals may associate with changes in age‐related immigration rates of sub‐adult males.
... At the intra-annual level, savannas are characterized by a wet season and a dry season and, in ecosystems without predators, adult herbivores rarely die during the wet season when food is abundant. However, the greater vegetation cover in the wet season may help ambush predators increase their hunting success, as they can approach their prey undetected (Funston et al. 2001;Hopcraft et al. 2005). Such predation is likely additive. ...
This chapter reviews the mechanisms underlying consumptive and non‐consumptive effects of predators on prey. It provides an overview of consumptive and non‐consumptive mechanisms, the current knowledge about their importance, and how they may interact. The chapter draws from the theoretical literature and from field studies conducted in biomes other than savanna, focusing on studies involving large mammalian herbivores, when available, and then reviews the evidence that these mechanisms operate in savannas. The best evidence of consumptive effects of predation on herbivore populations is revealed when predators are either reintroduced or removed. In ecosystems where predators are present at ecologically effective densities, what is of importance for an herbivore population is whether the mortality due to predation is additive or compensatory. The chapter emphasizes the role of ecosystem productivity, landscape composition, and the composition and functioning of the carnivore community underlying these differences.
Within the realm of social carnivores, lions (Panthera leo) are the sole representative of the large felids. While other Felidae may tolerate temporary associations with dependent offspring or receptive members of the opposite sex, lions are the only big cat where both males and females live in permanent social units. Each sex forms independent groups that come together to engage in a wide variety of social behaviors, including cooperative hunting, mutual defense of kills, cooperative territorial defense, and communal raising of young. Unique patterns of egalitarianism and flexible fission-fusion dynamics make this social structure distinctive among mammals. The bulk of our knowledge on the evolutionary drivers of lion sociality has been derived from extensive studies of populations inhabiting mesic and productive eastern African savanna systems. However, lions across Africa occupy a wide variety of habitat types, including arid deserts, seasonally flooded wetlands, and densely thicketed scrub. Comparison of lion behavior and interactions across their range reveals that their social strategies are highly plastic and adapt to maximize survival and fitness under prevailing local conditions. In this chapter, we first review the ultimate drivers of lion gregariousness based on research from eastern Africa and discuss how lions optimize individual fitness under constraints imposed by group-living. We then explore how variation in proximate drivers of social living (i.e., resource availability, intensity of inter- and intra-specific competition, and habitat structure) can shape the expression of social behavior. We end with a discussion of the social adaptations lions make to survive in increasingly human-dominated landscapes.
King of the Beasts! Captured in legends of ancient Rome, Greece and Egypt and threading through traditions of African culture, its powerful spell created a symbol of royal leadership. It gave form to the Sphinx and engraved on Egyptian tombs. The eye and the mane gave rise to religious myths and beliefs in traditional African societies. One of the “big five” and great tourist and hunter attraction.
Explianed in the article the biological parameters of: Taxonomy, Distribution, Species Status, Description, Trophy, Habitat & Feeding, Social population order, Behaviour, Growth & Production, Live sale price history.
The lion’s name is derived from the Greek word “leon”; first described as Felis leo by Linnaeus (1758) from a specimen in Algeria – changed in 1917 (accepted in 1985) to Panthera leo splitting the cat family into felinae (purring cats) and pantherinae (roaring cats, an elastic ligament in the hyoid apparatus allows movement of the larynx and roaring), with four species: lion Panthera leo (subspecies P.l. leo the African lion, and P.l. persica the Asiatic lion), leopard P. pardus (3 sub-species), jaguar P. onca (America), and tiger P. tigris (Asia, 5 sub-species).
Earliest lion fossils date back 3,5 million years (ma) at Leatoli in Tanzania, and in South Africa at Sterkfontein caves ( dating 2.8-2.4 ma). Despite the genetic hybridization that gives rise to varieties such as the white lion of the Timbavati, only two living extant lion subspecies are recognized.
Distribution & Species status
Historically, lions occurred throughout Europe, Asia and Africa; became extinct in Europe c.1100s, and extinct in North Africa between 1890-1940, in 1990 <300 in Asia, and in 2004 <800 in South Africa. Global numbers declined from c.200 000 in 1880 to 20 000 in 2001, currently around 23 000, of which in South Africa c.3 500 in protected parks and >8 000 on private farms. The Kruger Park population all infested with tuberculosis. Lions disappeared from the Cape flats, Western, Northern and Eastern Cape and the greater of KwaZulu-Natal during the 1860s – 164 lions in Hluhluwe-Imfolozi in 1988 declined to 65 by 1992. Illegal (uncontrolled) black-market trade in lion bones to East–Southeast Asia has been noted as a major threat to lion species survival – controlled sustainable legal trade needs much expanded development. ......
Optimal foraging and landscape of fear theories provide frameworks which can be useful for investigating animal's space and prey use decisions. Predators, such as African lions Panthera leo, are likely to respond to prey abundance, accessibility, profitability and potential risks, often anthropogenic in nature, while making foraging decision. Identifying the relative role of these processes has important conservation implications. We investigated the relative role of responses to a pastoralist landscape of fear within lion feeding and spatial ecology in a landscape at the human-wildlands interface. We collected spatial and predation data from 12 GPS-collared lions and ungulate count data from transects, along the South Africa-Mozambique border, including parts of Kruger, Limpopo and Banhine National Parks. We calculated Jacobs' Index values from 80 kills to investigate lion selection of wild and domestic ungulates as prey, used maximum entropy modelling to predict multi-season ungulate spatial occurrence and used resource selection functions to estimate the relative probability of use of wild and domestic ungulate areas by lions. All lions had access to wild prey and domestic livestock within their home ranges. Lions showed a strong selection for large-bodied wild ungulates as prey taking waterbuck, zebra, kudu and buffalo more frequently then predicted by their availability. Lions showed a slight avoidance of cattle as prey, with cattle outnumbering larger ungulates across much of the study area. Lions showed the greatest selection for habitats with high occurrences of wild prey, specifically areas with kudu, then nyala and buffalo, during the dry seasons and showed strong avoidance of cattle areas during the wet season; a season when cattle are kept closer to settlements and thus better protected and easier to predict and avoid. These results suggest that lions select for wild prey and habitats optimally, yet show a fear response to cattle and cattle areas. This duality in the foraging behaviour of lions suggests that efforts to mitigate human-lion conflict and preserve vulnerable lion populations should focus on both increasing wild ungulate populations as well as exploiting lion's fear of humans with careful consideration of the risks of livestock presence acting as an ecological trap for vulnerable lion populations. K E Y W O R D S African lion, carnivore conservation, habitat use, human-wildlife conflict, landscape of fear, optimal foraging, prey selection, resource-selection function 13652028, 0, Downloaded from https://onlinelibrary.wiley.com
Olfaction is a key sense, enabling animals to locate forage, select mates, navigate their environment, and avoid predation. Wind is an important abiotic factor that modulates the strength of olfactory information detected by animals. In theory, when airflow is unidirectional, an animal can increase odor detection probability and maximize the amount of olfactory information gained by moving crosswind. Given energetic costs inherent to activity and locomotion, behavioral search strategies that optimize the benefit-cost ratio should be advantageous. We tested whether African lions (Panthera leo) modify their movement directionality and distance according to wind speed and direction during hours of darkness when they are most active. We tracked 29 lions in southern Zimbabwe using GPS collars and deployed a weather station to collect detailed abiotic data. We found that when wind speeds increased lions were more likely to move crosswind. We also found that female lions, which tend to hunt more often than males, traveled farther when wind speeds were stronger. The results of our analysis suggest that lions adjust their movement behavior according to wind speed and direction. We inferred that this was a behavioral decision to maximize the amount of olfactory information gained per unit of energy spent. Our findings not only offer one of the first detailed insights on large carnivore anemotaxis (movement direction relative to wind) but also make an important contribution towards understanding the influence of wind on predator ecology in general which remains understudied to date.
Introduced predators are one of the leading causes of decline in island vertebrates. Understanding how they hunt and kill threatened prey can help improve management activities. Although broadscale features are known to influence predator movement patterns, factors influencing fine scale movement are often overlooked. In particular, the influence of prey cues and microhabitat features has received little attention despite predators spending considerable time hunting prey using a range of visual, olfactory and auditory cues. Using feral cats as a case study, we used video and GPS collars combined with ground-truthing to determine if predators use fine-scale prey cues or microhabitat features to hunt in an arid environment.
Feral cat activity was comprised of continuous traverses interspersed with periods of stationary activity (GPS clusters) generally less than 40 minutes in duration. Video collars confirmed that these clusters included the majority of stalk and pounce hunting bouts. Stationary activity was significantly focussed on prominent prey cues such as burrows, foraging digs or warrens of mammalian prey including both exotic (rabbits) and threatened native species (rodents, bilbies and bettongs). Evidence of prey kills was higher at cluster sites. Cats spent significantly more time at microsites with high vegetation cover including single trees, suggesting that they use prominent prey cues and patches of thick cover to increase their probability of encountering prey and/or to conceal themselves during hunting or feeding activity. Results suggest prey species with conspicuous cues are at higher risk of predation and this vulnerability could increase over time as resident cats learn to identify the location of prey cues within their home range. Conversely, removing resident knowledgeable predators may reduce predation rates if immigrating predators take time to learn to locate prey cues in their new environment. We urge researchers to investigate fine-scale drivers of movement patterns as this information is likely to be critical for long term management of predator species.
Resource partitioning promotes coexistence among guild members, and carnivores reduce interference competition through behavioural mechanisms that promote spatio-temporal separation. We analyzed sympatric lion and spotted hyena movements and activity patterns to ascertain the mechanisms facilitating their coexistence within semi-arid and wetland ecosystems. We identified recurrent high-use (revisitation) and extended stay (duration) areas within home ranges, and correlated environmental variables with movement-derived measures of inter- and intraspecific interactions. Spatial overlaps among lions and hyenas occurred at edges of home ranges, around water-points, along pathways between patches of high-use areas, and expanded during the wet season. Lions shared more of their home ranges with spotted hyenas in arid ecosystems, but shared more of their ranges with conspecifics in mesic environments. Despite shared space use, we found evidence for subtle temporal differences in the nocturnal movement and activity patterns between the two predators, suggesting a fine localized-scale avoidance strategy. Revisitation frequency and duration within home ranges were influenced by interspecific interactions, subsequent to land cover categories and diel cycles. Intraspecific interactions were also important for lions and, for hyenas, moon illumination and ungulates attracted to former anthrax carcass sites in Etosha, with distance to water in Chobe/Linyanti. Recursion and duration according to locales of competitor probabilities were similar among female lions and both sexes of hyenas, but different for male lions. Our results suggest that lions and spotted hyenas mediate the potential for interference competition through subtle differences in temporal activity, fine-scale habitat use differentiation, and localized reactive-avoidance behaviours. These findings enhance our understanding of the potential effects of interspecific interactions among large carnivore space-use patterns within an apex predator system, and show adaptability across heterogeneous and homogeneous environments. Future conservation plans should emphasize the importance of inter- and intraspecific competition within large carnivore communities, particularly moderating such effects within increasingly fragmented landscapes.
The large felid carnivores are among the most endangered, and the most challenging, species to conserve on this increasingly human-dominated planet. In modern times, large felid carnivores were widely distributed in the continents of Asia, Africa, and the Americas. Unfortunately, global human expansion, loss of prey species, hunting and poaching, and retaliatory killings after livestock predation have greatly reduced and fragmented their original ranges and decimated their populations. In this chapter, large felid carnivore characteristics, usual habitats, ecology, and predatory behaviors are reviewed. Changes in great cat distribution, changes in wild prey populations resulting in a shift to increased livestock predation, and the resulting human-felid conflicts are discussed. Preservation of remaining wild large felid carnivore populations has become a global conservation priority as populations have plummeted over the last century. Current approaches to better understand and conserve these apex keystone predators and to maintain ecosystem integrity are discussed. Current strategies and policies to ameliorate and resolve the intricate and difficult problems of predator-human conflict are examined. The complex issues of “problem carnivores” and “man-eaters” are discussed. Finally, recommendations on creative, fluid, and scientifically sound strategies that might be employed to address these conflicts in a manner acceptable to all key stakeholders are discussed.
Two Panthera leo group sizes maximize foraging success during the season of prey scarcity: one female and 5-6 females. Foraging success does not vary significantly with group size when prey is abundant. Female lions live in fission-fusion social units (prides) and forage only with members of their own pride. If lion grouping patterns were primarily related to group-size-specific feeding efficiency, females in prides containing <5 females should forage alone when prey is scarce; females in larger prides should forage alone or in groups of 5-6. However, females in small prides most commonly forage in as large a group as possible, even at the expense of foraging efficiency. Females in large prides most often forage in intermediate group sizes of 4 or 5. However, mothers keep their cubs in a creche and form highly stable maternity groups that are effective in defending the cubs against infanticidal males. Most large prides contain a creche involving 4 or 5 mothers; in the absence of a creche, large prides show no preference for any group size. Females also compete aggressively against neighboring prides, and larger groups successfully repel smaller ones in territorial disputes. Small prides appear to be excessively gregarious in order to compete against larger neighboring prides. -from Authors
Lion hunting behaviour was studied in two regions of Queen Elizabeth National Park, Uganda. Six environmental, three prey-related and two lion-related factors were evaluated for their influence on hunting success. Grass and bush cover, hunting group size, prey group size and the presence/absence of the moon (on nocturnal hunts) had significant effect on the outcome of hunts. Interspecific differences in escape methods used by prey are also discussed.Cover, prey availability and prey body size appeared to be the major causes of variation in lion foraging behaviour between study sites. In Mweya, lions hunted opportunistically for small prey species during the day, while at night, the pride actively searched for large prey. In Ishasha, lions foraged most often during moonless nights, when their hunting success was greatest.RésuméLe comportement de chasse du lion fut étudié dans deux régions du Queen Elizabeth National Park en Ouganda. L'influence de six facteurs environnementaux, trois facteurs liés aux proies et deux facteurs liés au lion lui-même furent évalués dans le cadre du succès de la chasse. La couverture des graminées et des buissons, la taille du groupe en chasse, la taille du groupe des proies et la présence/absence de la lune (durant les chasses nocturnes) ont un effet significatif sur la conclusion des chasses. Les différences interspécifiques des méthodes de fuite utilisées par la proie sont aussi discutées.Le couvert végétal, la disponibilité de proies et la taille corporelle de la proie paraissent être les causes majeures de variation du comportement de recherche du lion entre les sites d'étude. A Mweya, des lions chassent de manière opportuniste des espèces de proie de petite taille durant la journée, alors que de nuit, la bande cherche activement des grandes proie. A Ashasha, des lions chassent le plus souvent durant les nuits sans lune, lorsque leurs chances de succès sont les plus élevées.
Two methods of estimating the number of lions in the Kalahari Gemsbok National Park, Republic of South Africa, are described; the first gives a minimum figure (113) and the second a more realistic one (140). Data are presented on sex and age ratios and pride composition. The factors contributing to the low density are briefly discussed and some management practices in connection with lions trespassing out of the Park are suggested.
Knowledge on the abiotic and biotic components of the Kruger National Park (KNP) system has increased to such an extent, that it was possible to zonate the KNP into landscapes. A landscape was defined as an area with a specific geomorphology, climate, soil and vegetation pattern together with the associated fauna. On this basis 35 landscapes were identified and described in terms of the components mentioned in the definition. The objective of classification is that future management should be based on these landscapes. Relevant management considerations may change, but the landscape a@ a basic functional unit should not be negotiable.
Five methods for studying food habits of large African carnivores are evaluated. Fecal analysis is useful for a basic description of the diet, provided that an adequate sample of scats can be obtained. However, it is impossible to quantify the amount eaten and to determine the proportions of killed versus scavenged food. Tracking spoor in restricted habitats is useful for most species, except the brown hyena (Hyaena brunnea). Opportunistic, and for most species, radio-location observations are biased towards large prey animals, because small animals are eaten quickly, leaving no trace. However, the data can be used to study sex and age selection of adult prey. Direct observations provide accurate measurements of consumption rates, killing frequency, and prey selection, provided they can be carried out without disturbing predator or prey. Mixing data from incompatible techniques must be avoided.
The role of Panthera leo predation in the dynamics of blue wildebeest Connochaetes taurinus and zebra Equus burchelli populations was investigated through simulation models. Data used in the models were from observations in SE Kruger National Park. Model 1 ascertained the number of killing lions (adult females) that could be supported by each prey population while remaining stable. A single model was constructed for the sedentary wildebeest population. A summer and winter model was constructed for the semi-migratory zebra population. The sensitivity of the parameters in the model was tested by changing their value by 10%. In model 2, the kill age structure for each species was changed to determine the number of killing lions the altered prey selection parameters could support. There was no difference in the vulnerability of either species to predation. Zebra foals (<1 yr) were killed more frequently than expected. No selection for sex or by season could be found for either species. Model 1 predicted that the wildebeest population stabilizes with 7.7 killing lions, close to the number in the study area. The winter zebra populations stabilizes with 6.8 killing lions and the summer zebra population with 19.4. Manipulation of kill rate followed by adult fecundity rate had the greatest effect on population size of both species. In model 2, wildebeest predation was made selective towards calves and zebra predation was made non-selective for sex and age. With these parameters the wildebeest population stabilizes with 10.7 killing lions and the zebra population with 5.4 in winter and 15.1 in summer. The models suggest that lion predation affected wildebeest more severely than zebra during the study. This was through the way in which lions selected their prey, and because of the sedentary behaviour of the wildebeest, as opposed to the semi-migratory behaviour of the zebra. -from Authors
The effect of the time of exposure to an aerosol of viable microorganisms on the incidence of respiratory infections associated with a 3 h exposure to ozone (O3) was studied. The 157 and 196 (0.08–0.1 ppm) levels of O3 used occur regularly in some urban communities. The studies reported here show that the susceptibility of mice to a laboratory-induced infection can be maximally enhanced if the infectious challenge is concurrent with the exposure to O3.
1. Aspects of the influence of predation on the African buffalo Syncerus caffer were studied in Lake Manyara National Park, northern Tanzania, from 1981 to 1985. 2. During the study period, predation by lions was the most important cause of death in buffalo but occasionally major epidemics, such as rinderpest, can be by far the most important cause of death. 3. A comparison of Manyara with four other areas in Africa shows that the Manyara buffalo ran the same level of risk as those in these other areas, and that in periods in which epidemics or drought conditions are absent it was normal that ultimately some 90% of the large herbivores fell victim to predation. 4. Juvenile mortality in females was higher than in males but adult mortality in females was lower than in males. Adult bulls ran a much higher risk to be killed by lions (12.9% p.a.) than herd-living buffalo (4.4% p.a.); the latter category comprised cows and sub-adult bulls. 5. Risk of predation was highest near the ecotone between grasslands and structurally closed vegetations. Grasslands were as dangerous as woodlands and thickets, and only the mudflats were really safe. The chance to be killed fell steeply away from (potential) cover for lions. 6. Buffalo were more vigilant at night than during the day, peripheral animals more so than central ones. Adult bulls were more vigilant than other categories of buffalo. It is concluded that vigilance in buffalo was not restricted to predator detection. 7. The choice of feeding areas in buffalo was not dictated by predation risk alone.
Direct observations on the foraging activities of lions, based on 920 attempts to capture prey and 156 kills, on the plains of Etosha National Park, Namibia, revealed a regular nocturnal pattern of mostly coordinated group hunting. Lions scavenged rarely and killed mainly prey animals weighing less than 50 kg, which contributed to 73% of the observed kills and 50% of the estimated biomass consumed. Capture success increased with lion group size and was also greater during coordinated group hunts. Lions hunted most of the prey that they encountered, showing a preference for large prey species. Average food acquisition ranged from 8.7 kg/day per lioness in the dry season to 14 kg/day per lioness in the wet season. During the dry season, coordinated cooperative hunting was essential and lionesses most often formed groups of 2, thereby acquiring higher daily food intake than groups of other sizes. In the wet season, lioness groups of all sizes obtained more than the estimated daily requirements, and lionesses did not uniformly forage in the smaller groups capable of greater food acquisition.
The capture of prey by the African lion in Ngorongoro Crater, Tanzania, is examined. For analytic purposes the process is envisaged as a consecutive set of events: searching, stalking, attacking, and subduing prey.Initiation of prey capture activity by the lion arises from the interaction of degree of stimulation presented by the prey, timing of the activity cycle of the lion (diurnal or nocturnal), and its level of hunger. These factors yield thresholds for initiation which result in kills being directly related to prey density at high and low densities and inversely density dependent at intermediate prey densities.Success in stalking appears to depend primarily upon the failure of the prey to see the approaching lion until it is within its effective distance. Using data from actual stalks and contrived experiments, the probability is calculated of the lion being detected for a range of conditions.The attack involves a matching of the sprinting abilities of the predator and prey. To analyze this aspect in detail, the velocity curves of running are defined for the lion and four prey species.
Individually identified lions (Panthera leo) were observed on the open, semi-arid plains in Namibia. Data from 486 coordinated group hunts were analysed to assess cooperation and individual variation in hunting tactics. Group hunts generally involved a formation whereby some lionesses (wings) circled prey while others (centres) waited for prey to move towards them. Those lionesses that occupied wing stalking roles frequently initiated an attack on the prey, while lionesses in centre roles moved relatively small distances and most often captured prey in flight from other lionesses. Each lioness in a given pride repeatedly occupied the same position in a hunting formation. Hunts where most lionesses present occupied their preferred positions had a high probability of success. Individual hunting behaviour was not inflexible, however, but varied according to different group compositions and to variations in the behaviour of other individuals present. The role of cooperative hunting and its apparent advantages within the semi-arid environment of Etosha National Park, Namibia, are discussed.
The participation of individual African lions, Panthera leo, during 64 communal hunts of four prey species was measured to quantify the extent to which lions cooperate and the factors affecting the degree of cooperation. The extent of individual participation in communal hunts varied significantly. Finite mixture models were used to determine the probability that each lion's behaviour belonged to each of three strategies: ‘refraining’ (non-participation in hunts), ‘conforming’ (active participation in groups in which all individuals behaved similarly) and ‘pursuing’ (active participation in groups where individual behaviour varies). Our analysis reveals two important trends. First, males refrain more and pursue less than females. Second, refraining during a group hunt is more common during hunts of prey that appear to be easier to capture: lions are more likely to refrain during hunts of wart hog, Phacochoerus aethiopicus, and less likely to refrain during hunts of zebra, Equus burchelli, and buffalo, Syncerus caffer. Of the alternatives considered, the data indicate that refraining is ‘cheating’ and that lions exhibiting this strategy are thus exploiting the hunting behaviour of their companions. These results are discussed in the framework of a recent game-theoretical model of cooperative hunting.
In the Kruger National Park, male lions, Panthera leo, acquire most of their food by hunting rather than scavenging. This study, the most intensive to date of male lion ecology, showed that in savanna woodlands, with high buffalo, Syncerus caffer, densities, male lions were frequent and successful hunters. The main prey species of all male group types, but particularly nonterritorial males, was buffalo. By contrast, females preyed more frequently on the most abundant medium-sized ungulates, such as wildebeest, Connochaetes taurinus, and zebra, Equus burchelli. Thus intraspecific prey selection separation was based primarily on intersexual and, to a lesser extent, social differences. Furthermore, both nonterritorial males and pride females located their favoured prey, buffalo and medium-sized ungulates, respectively, more often than other prey. We investigated the influence of several ecological variables on the socioecology of male lions, particularly as we had determined that territorial males spent little time with their pride females and tended to hunt by themselves in their respective male coalitions. Further analysis showed that in a range of ecosystems in southern and eastern Africa the proportion of time territorial males spent with, and thus scavenged from, their pride females was strongly influenced by vegetation structure, and therefore probably by the assemblage of available ungulates. In open systems, territorial males were, therefore, likely to be encountered with pride females, whereas in more wooded areas they were likely to be encountered away from their pride females. We suggest that this is because vegetation structure influences food/prey availability and hunting success and influences territory maintenance and/or cub defence. (c) 1998 The Association for the Study of Animal Behaviour.
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