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Food caching is a common strategy used by a diversity of animals, including carnivores, to store and/or secure food. Despite its prevalence, the drivers of caching behaviour, and its impacts on individuals, remain poorly understood, particularly for short-term food cachers. Leopards Panthera pardus exhibit a unique form of short-term food caching, regularly hoisting, storing, and consuming prey in trees. We explored the factors motivating such behaviour among leopards in the Sabi Sand Game Reserve, South Africa, associated with four not mutually exclusive hypotheses: food-perishability, consumption-time, resource-pulse, and kleptoparasitism-avoidance. Using data from 2,032 prey items killed by 104 leopards from 2013–2015, we built generalized linear mixed models to examine how hoisting behaviour, feeding time, and the likelihood of a kill being kleptoparasitised varied with leopard sex and age, prey size and vulnerability, vegetation, elevation, climate, and the short-term (immediate presence) and long-term (species density) risk posed by dominant competitors. Leopards hoisted 51% of kills. They were more likely to hoist kills of an intermediate size, outside of a resource pulse, and in response to the presence of some competitors. Hoisted kills were also fed on for longer than non-hoisted kills. At least 21% of kills were kleptoparasitised, mainly by spotted hyaenas Crocuta crocuta. Kills were more likely to be kleptoparasitised at lower temperatures and if prey were larger, not hoisted, and in areas where the risk of encountering hyaenas was greatest. Female leopards that suffered higher rates of kleptoparasitism exhibited lower annual reproductive success than females that lost fewer kills. Our results strongly support the kleptoparasitism-avoidance hypothesis and suggest hoisting is a key adaptation that enables leopards to coexist sympatrically with high densities of competitors. We further argue that leopards may select smaller-sized prey than predicted by optimal foraging theory, to balance trade-offs between kleptoparasitic losses and the energetic gains derived from killing larger prey. Although caching may provide the added benefits of delaying food perishability and enabling consumption over an extended period, the behaviour primarily appears to be a strategy for leopards, and possibly other short-term cachers, to reduce the risks of kleptoparasitism. This article is protected by copyright. All rights reserved.

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... In one study, lions succeeded in stealing hyena kills with every attempt, even when outnumbered six to one [113]. Leopards in some populations frequently store and consume prey in trees [114,115], a unique strategy believed to be employed to avoid kleptoparasitism. A study in a South African reserve identified a 21% steal rate of leopard prey, with the primary kleptoparasite being the spotted hyena [115]. ...
... Leopards in some populations frequently store and consume prey in trees [114,115], a unique strategy believed to be employed to avoid kleptoparasitism. A study in a South African reserve identified a 21% steal rate of leopard prey, with the primary kleptoparasite being the spotted hyena [115]. Brown hyenas are unique in the large African carnivore guild as in many regions they are primarily scavengers. ...
... Lions are capable of stealing kills from Africa's top predators, including spotted hyenas [113] and leopards [115]. Loarie et al. [104] assessed the effect of vegetation structure on lion predation in KNP using Light Detection and Ranging (LiDAR) and GPS-tracking of lions. ...
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Bush encroachment is a habitat change phenomenon that threatens savanna and grassland ecosystems worldwide. In Africa, large carnivores in bush encroached landscapes must adjust to increasing woody plant cover and biomass, which could affect predation success at multiple stages through complex and context-dependent pathways. We highlight, interpret, and compare studies that assessed how bush encroachment or related habitat parameters affect the predation stages of large African carnivores. Bush encroachment may directly or indirectly affect predation success in various ways, including by: (1) altering habitat structure, which may affect hunting efficiency and prey accessibility; (2) changing prey abundance/distribution, with smaller species and browsers being potentially favoured; (3) influencing interference competition within the carnivore guild. For habitat or dietary specialists, and subordinate predators that are vulnerable to both top-down and bottom-up ecosystem effects, these alterations may be detrimental and eventually incur population fitness costs. As the threat of bush encroachment continues, future studies are required to assess indirect effects on competitive interactions within the large African carnivore guild to ensure that conservation efforts are focused. Additionally, to better understand the effects of bush encroachment across Africa, further research is necessary in affected areas as overall little attention has been devoted to the topic.
... On average, 98 AE 2 vehicles are active throughout the SSGR per game drive, resulting in a mean density of one vehicle per 6 km 2 . The high number of vehicles, together with an extensive road network (a total length of 3159 km and mean density of five roads per km 2 ), ensures that most of the reserve is covered daily (Balme, Miller, et al., 2017). Game drives are also not limited to roads; a skilled tracker on the front of each vehicle scans for signs of charismatic species such as leopards. ...
... Due to this intensive search effort and the relaxed nature of individuals, leopard sightings are frequent; on average, 6967 AE 377 unique leopard sightings are recorded each year. The accuracy of individual identification of leopards in SSGR was previously assessed using photographs of known individuals and identification by guides was correct in all instances (Balme, Miller, et al., 2017;Balme, Pitman, et al., 2017). Similarly, cross-referencing records of the same sighting from different lodges yielded no significant discrepancies . ...
... Due to this intensive search effort and the relaxed nature of individuals, leopard sightings are frequent; on average, 6967 AE 377 unique leopard sightings are recorded each year. The accuracy of individual identification of leopards in SSGR was previously assessed using photographs of known individuals and identification by guides was correct in all instances (Balme, Miller, et al., 2017;Balme, Pitman, et al., 2017). Similarly, cross-referencing records of the same sighting from different lodges yielded no significant discrepancies . ...
Article
Variation in home range size exists among and within wildlife populations. Home range size variation may be driven by both intrinsic and extrinsic factors, including sex, food and reproductive resources, density and competition. In this study, we investigated the sex‐specific impacts of prey and reproductive resources, conspecific density and competition on leopard Panthera pardus home range size at two spatio‐temporal scales in the Sabi Sand Game Reserve, South Africa. Male leopard home ranges were more than twice the size of those of females, in line with expectations for a solitary, polygamous species. Both male and female leopard space‐use were primarily driven by short‐term changes in intra‐sexual conspecific density. Females were influenced by both short and long‐term drivers, with long‐term prey availability (home range and core) and refugia (core) further impacting size. Males were almost exclusively influenced by short‐term drivers; home range size was further impacted by short‐term changes in female leopard and prey density, and age. Long‐term prey availability contributed to male leopard core size. The difference in impact of short‐ and long‐term drivers between the sexes likely relates to tenure expectations; males may be forced out of their territories at any time and should therefore optimize their space‐use based on present conditions. Female leopards, however, must secure a home range that maximizes their reproductive success in the short‐ and long‐term in order to raise cubs to independence. Our findings challenge expectations that space‐use is primarily resource‐driven and demonstrate the critical role of social factors in saturated populations of solitary species. Furthermore, we illustrate the importance of considering temporally variable factors across different timescales to fully understand their impact on mammalian spatial organization. We investigated the sex‐specific impacts of prey and reproductive resources, conspecific density and competition on leopard Panthera pardus home range size at two spatio‐temporal scales. Male home ranges were double the size of female home ranges, and both male and female leopard home range size were primarily driven by short‐term changes in intra‐sexual conspecific density. Females were influenced by short‐ and long‐term patterns, while males were influenced almost exclusively by short‐term changes; this is likely linked to the disproportionate investment in reproduction between the sexes. Our findings challenge expectations that space‐use is primarily resource‐driven and demonstrate the complex interplay of ecological and social factors in natural populations of solitary species
... Food caching, defined as storing and/or securing food, is an evolutionary strategy adopted by predators which can reduce kleptoparasitism or safeguard surplus food for future consumption. It is common in carnivores from red foxes Vulpes vulpes [19] to leopards Panthera pardus [20] and grizzly bears Ursus arctos [21]. Identifying when food is cached is useful for characterising kill sites, and consequently quantifying predator-prey dynamics [22,23]. ...
... Importantly, in the absence of other large carnivores, kleptoparasitism by conspecifics can still be a major source of losses [20]. The high population density (5.6 individuals/100 km 2 [47]) and large home range overlap between neighboring conspecifics in Tandoureh [27] make caching behaviour important for avoiding this [20,48]. ...
... Importantly, in the absence of other large carnivores, kleptoparasitism by conspecifics can still be a major source of losses [20]. The high population density (5.6 individuals/100 km 2 [47]) and large home range overlap between neighboring conspecifics in Tandoureh [27] make caching behaviour important for avoiding this [20,48]. ...
Article
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Background Tackling behavioural questions often requires identifying points in space and time where animals make decisions and linking these to environmental variables. State-space modeling is useful for analysing movement trajectories, particularly with hidden Markov models (HMM). Yet importantly, the ontogeny of underlying (unobservable) behavioural states revealed by the HMMs has rarely been verified in the field. Methods Using hidden Markov models of individual movement from animal location, biotelemetry, and environmental data, we explored multistate behaviour and the effect of associated intrinsic and extrinsic drivers across life stages. We also decomposed the activity budgets of different movement states at two general and caching phases. The latter - defined as the period following a kill which likely involves the caching of uneaten prey - was subsequently confirmed by field inspections. We applied this method to GPS relocation data of a caching predator, Persian leopard Panthera pardus saxicolor in northeastern Iran. Results Multistate modeling provided strong evidence for an effect of life stage on the behavioural states and their associated time budget. Although environmental covariates (ambient temperature and diel period) and ecological outcomes (predation) affected behavioural states in non-resident leopards, the response in resident leopards was not clear, except that temporal patterns were consistent with a crepuscular and nocturnal movement pattern. Resident leopards adopt an energetically more costly mobile behaviour for most of their time while non-residents shift their behavioural states from high energetic expenditure states to energetically less costly encamped behaviour for most of their time, which is likely to be a risk avoidance strategy against conspecifics or humans. Conclusions This study demonstrates that plasticity in predator behaviour depending on life stage may tackle a trade-off between successful predation and avoiding the risks associated with conspecifics, human presence and maintaining home range. Range residency in territorial predators is energetically demanding and can outweigh the predator’s response to intrinsic and extrinsic variables such as thermoregulation or foraging needs. Our approach provides an insight into spatial behavior and decision making of leopards, and other large felids in rugged landscapes through the application of the HMMs in movement ecology.
... Foraging theory predicts that an apex predator should prefer prey of sufficient mass to be the most energy efficient, while also posing the least risk of injury to the predator (Stephens & Krebs, 1986). Additionally, for large carnivores that are not the apex of a carnivore intraguild, risk of kleptoparasitism is also a factor affecting prey preference and interactions (Macdonald, 1976;Balme et al., 2017). Means of reducing kleptoparasitism between large carnivores could thus be through niche differentiation (Balme, Hunter & Slotow, 2007), caching prey so that it remains out of reach (Balme et al., 2017), and temporal partitioning (Hayward & Slotow, 2009). ...
... Additionally, for large carnivores that are not the apex of a carnivore intraguild, risk of kleptoparasitism is also a factor affecting prey preference and interactions (Macdonald, 1976;Balme et al., 2017). Means of reducing kleptoparasitism between large carnivores could thus be through niche differentiation (Balme, Hunter & Slotow, 2007), caching prey so that it remains out of reach (Balme et al., 2017), and temporal partitioning (Hayward & Slotow, 2009). ...
... Leopards (Panthera pardus) have been found to exhibit more diurnal activity patterns in rainforests of Gabon (Henschel & Ray, 2003) and Thailand (Ngoprasert, Lynam & Gale, 2007), but more often exhibit cathemeral activity patterns in African savannah landscapes (Hayward & Slotow, 2009). A possible explanation for this has been to minimize kleptoparasitism and competition with lions (Panthera leo) and spotted hyenas (Crocuta crocuta), which are absent in rainforests (Hayward & Kerley, 2008;Hayward & Slotow, 2009;Balme et al., 2017). ...
Article
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Studying activity patterns and temporal overlap among carnivores and their putative prey is difficult because of their secretive and elusive nature. With large carnivores declining worldwide, it is imperative for conservation planning that we understand how large carnivores interact with their prey and competitors. Camera trapping offers a promising avenue to address this issue. We investigated temporal overlap between male and female leopards, their known and putative prey as well as their competitor, the spotted hyenas, in the Udzungwa Mountains of Tanzania. Data consisted of 4297 independent events from a 30 min interval criterion from 164 camera trap sites we sampled. Leopards were captured by camera traps throughout the day, with male and female leopards showing significantly different activity patterns (P < 0.001) indicating sexual segregation in activity patterns, with male leopards being more nocturnal than female leopards. Leopards had significantly different activity patterns from that of the majority of their prey, with yellow baboons, that displayed peak activity during midday, that had the least overlap. Moreover, both male and female leopards had significantly different activity patterns from that of spotted hyenas (P = <0.001), with female leopards appearing to be inactive during hours with peak hyena activity. We conclude that systematic camera trapping is a useful tool to study activity patterns and temporal niche interactions between sym-patric carnivores and, to a lesser extent, their prey.
... Secondly, challenges in capturing spotted and brown hyaenas led to their omission from our study. However, spotted hyaenas can impact the reproductive fitness of leopards through kleptoparasitism (see Balme et al., 2017a). Consequently, given the high density of spotted hyaena within our study area and their impact on leopards elsewhere (Balme et al., 2017a), it is possible that they exert a top-down effect on leopard spatiotemporal patterns that we were unable to measure in this study. ...
... However, spotted hyaenas can impact the reproductive fitness of leopards through kleptoparasitism (see Balme et al., 2017a). Consequently, given the high density of spotted hyaena within our study area and their impact on leopards elsewhere (Balme et al., 2017a), it is possible that they exert a top-down effect on leopard spatiotemporal patterns that we were unable to measure in this study. ...
... That said, in the absence of large scale spatial or temporal avoidance of competitors, it is likely that leopards adapt finer-scale behaviours, such as dragging prey into trees, to perceived competitor risk or during the occurrence of direct competitor encounters. However, beyond prey caching (Stein et al., 2015;Balme et al., 2017a), little is known of these coexistence mechanisms. Whilst such interactions were beyond the scope of this study, investigations into fine-scale coexistence mechanisms will provide greater insights into the factors allowing leopards to coexist within such a highly competitive predator assemblage. ...
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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. 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). We used overlapping spatio‐temporal data from African predators to examine spatial interactions and temporal partitioning between leopards and other guild members in northern Botswana. Our results suggest predator avoidance has a limited impact on broad‐scale leopard spatial and temporal niches in our study area.
... Leopards tend to display some degree of dietary overlap with their major competitors (du Preez, Purdon, Trethowan, MacDonald, & Loveridge, 2017;Hayward & Kerley, 2008;Mbizah, Marino, & Groom, 2012), but they are vulnerable to exploitative competition. For example, a recent study found that 21% of leopard kills were kleptoparasitised, mostly by hyaenas, while lions posed a direct threat to leopard survival (Balme, Miller, Pitman, & Hunter, 2017). However, leopards have a remarkable adaptability to intraguild competition, both in terms of their behaviour and diet (Karanth & Sunquist, 1995). ...
... However, leopards have a remarkable adaptability to intraguild competition, both in terms of their behaviour and diet (Karanth & Sunquist, 1995). For example, previous studies in Africa have also shown that leopards may reduce potential conflict with other predators by specialising on specific prey (Voigt, Krofel, Menges, Wachter, & Melzheimer, 2018), selecting relatively smaller prey and hoisting/caching prey in trees away from more dominant competitors (Bailey, 1993;Balme et al., 2017;Stein, Bourquin, & McNutt, 2014). As a result, leopards are able to live in sympatry with other competitors and often thrive in these multicarnivore systems (Bailey, 1993;Hayward & Kerley, 2008). ...
... We recorded preference for bushbuck, common duiker, nyala and reedbuck in our study area, which coincided somewhat with the preferred prey found by Hayward et al. (2006). These species typically involve less risk when hunting are generally easier to subdue and hoist into trees compared to larger prey species and include greater energetic gains compared to considerably smaller prey (Balme et al., 2017;Hayward et al., 2006). Interestingly, Balme, Hunter, and Slotow (2007) (Mbizah et al., 2012). ...
Article
en 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. Résumé fr Entre 2011 et 2012, la guilde des carnivores de la réserve faunique de Majete (MWR), au Malawi, a été restaurée suite à la réintroduction du lion (Panthera leo) et du léopard (Panthera pardus). Le but de cette étude était de décrire et de comparer le régime alimentaire du lion, du léopard et de la hyène tachetée (Crocuta crocuta), déjà résidente, en analysant leurs excréments. Les lions et les hyènes présentaient le plus grand chevauchement alimentaire (O ab = 088) et sélectionnaient principalement des proies appartenant à des espèces de taille moyenne à grande. Les lions préféraient des proies dont le poids moyen était de 120.33 ± 42.14 kg (SE), avec le phacochère (Phacochoerus africanus) et le waterbuck (Kobus ellipsiprymnus) représentant 60.64% de la biomasse relative consommée. Les hyènes préféraient des proies dont le poids moyen était de 102.40 ± 41.69 kg et avaient une alimentation plus généralisée (B a = 0.46) par rapport aux lions (B a = 0.36). En revanche, les léopards occupaient une niche alimentaire sensiblement inférieure à celle des lions et des hyènes, sélectionnant des proies relativement plus petites dont le poids moyen était de 27.50 ± 6.74 kg. Nos résultats suggèrent que la coexistence entre la hyène (résidente), le lion et le léopard (tous deux réintroduits) dans MWR est facilitée par le partitionnement alimentaire. Nous conseillons un suivi à long terme lors de la réintroduction de carnivores dans de petites réserves clôturées, pour évaluer leur impact sur les populations de prédateurs et de proies.
... Leopard populations can also be suppressed by dominant carnivores, as documented in Asia for tigers (Panthera tigris) (Harihar et al. 2011), and suggested in Africa for African lions (Panthera leo), and spotted hyenas (Crocuta crocuta) (Packer et al. 2011). Besides interference competition, one of the possible mechanism of top-down suppression could be connected with kleptoparasitism, which was shown to reduce reproductive success in African leopards (Balme et al. 2017a). ...
... We did not test for the causation of the effects of dominant carnivores, but (Balme et al. 2019) found that leopard detections correlated with areas with high hyena detections was actually caused by hyenas seeking out and following leopards in order to kleptoparasitize their killed prey. Lions and hyenas kleptoparasitize leopards, which can have negative effects on leopard reproductive success (Balme et al. 2017a) and cause leopards in areas with higher abundance of dominant carnivores to feed on smaller prey (Hayward et al. 2006) and cache more of their prey in trees to avoid loss of food (Stein et al. 2015;Balme et al. 2017a). ...
... We did not test for the causation of the effects of dominant carnivores, but (Balme et al. 2019) found that leopard detections correlated with areas with high hyena detections was actually caused by hyenas seeking out and following leopards in order to kleptoparasitize their killed prey. Lions and hyenas kleptoparasitize leopards, which can have negative effects on leopard reproductive success (Balme et al. 2017a) and cause leopards in areas with higher abundance of dominant carnivores to feed on smaller prey (Hayward et al. 2006) and cache more of their prey in trees to avoid loss of food (Stein et al. 2015;Balme et al. 2017a). ...
Article
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Large carnivore conservation is important for ecosystem integrity and understanding drivers of their abundance is essential to guide conservation efforts. Leopard (Panthera pardus) populations are in a general state of decline, although local studies demonstrated large variation in their population trends and density estimates vary widely across their range. We used spatially-explicit capture-recapture models for unmarked populations with camera trap data from a citizen science project to estimate previously-unknown leopard population densities in Serengeti National Park, Tanzania, and determine potential biological drivers of their abundance and distribution. We estimated leopard densities, at 5.41 (95% CrI = 2.23–9.26) and 5.72 (95% CrI = 2.44–9.55) individuals/100 km2, in the dry and wet season, respectively, which confirmed Serengeti National Park as one of the strongholds of this species in Africa. In contrast to abundance estimates, we found that drivers of leopard abundance and distribution varied among the dry and wet seasons, and were primarily affected by interactions with other larger carnivores and cover. The underlying driver of leopard distribution may be the dynamic prey availability which shifts between seasons, leading to an avoidance of dominant carnivores when prey availability is low in the dry season but an association with dominant carnivores when prey availability is high in the wet season. As efforts to conserve large carnivore populations increase worldwide, our results highlight the benefits of using data from citizen science projects, including large camera-trapping surveys, to estimate local carnivore abundances. Using a Bayesian framework allows of estimation of population density, but it is also important to understand the factors that dictate their distribution across the year to inform conservation efforts.
... Interspecific competition between predators occurs when species exploit the same resources in space and time. This process can often manifest as kleptoparasitism, whereby one species steals resources such as food from the other, which can have pervasive effects from the individual to ecosystemlevel (Gorman et al. 1998, Krofel et al. 2012, Moleón et al. 2014, Balme et al. 2017). On the African continent, kleptoparasitism has been well documented among lions Panthera leo, leopards P. pardus, spotted hyaenas Crocuta crocuta, cheetahs Acinonyx jubatus, and African wild dogs Lycaon pictus (Scantlebury et al. 2014, Broekhuis and Irungu 2016, Balme et al. 2017). ...
... This process can often manifest as kleptoparasitism, whereby one species steals resources such as food from the other, which can have pervasive effects from the individual to ecosystemlevel (Gorman et al. 1998, Krofel et al. 2012, Moleón et al. 2014, Balme et al. 2017). On the African continent, kleptoparasitism has been well documented among lions Panthera leo, leopards P. pardus, spotted hyaenas Crocuta crocuta, cheetahs Acinonyx jubatus, and African wild dogs Lycaon pictus (Scantlebury et al. 2014, Broekhuis and Irungu 2016, Balme et al. 2017). However, little is known about this behaviour in medium-sized predators or the mesopredator guild (e.g. ...
... By contrast, prior observations of successful jackal scrounging associated with honey badgers exclusively involved small rodents (Muridae, Begg et al. (in press)). Kleptoparasitic interactions are often positively associated with prey size, with larger prey likely to be perceived by the kleptoparasite as representing a greater reward (Dies and Dies 2005, Dill and Davis 2012, Balme et al. 2017. While larger prey items represent a small part of the honey badger diet (Begg et al. 2003), they also represent prey items which are not commonly scrounged by black-backed jackals (Begg et al. in press). ...
Article
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Intra-guild competition, including kleptoparasitism, can shape a species’ ecology, particularly when competitors commonly occur sympatrically. Here, I describe a series of interspecific interactions between a black-backed jackal Canis mesomelas and two honey badgers Mellivora capensis from an observation in Etosha National Park, Namibia. This interaction involved a typical kleptoparasitic “producer-scrounger” relationship, but resulted in a novel observation of one honey badger successful kleptoparasitising the jackal to reclaim its prey. This observation highlights the complexity of foraging interactions between these two functionally important mesopredators and complements our current understanding of their foraging ecologies. In Canid Biology & Conservation (https://www.canids.org/c-b-c). Cite as: Gorta, S.B.Z. 2020. What goes around comes around: complex competitive interactions between two widespread southern African mesopredators. Canid Biology & Conservation 22(2):8-10. URL: http://www.canids.org/CBC/22/Interactions_between_African_mesopredators.pdf.
... Feeding in the presence of competing carnivores means leopards must balance food intake and risk (Brown, 1988;Verdolin, 2006). Leopards respond to kleptoparasitism by caching their kills in trees, which reduces interference from non-climbers such as hyenas, but not from competitors that are capable of climbing such as conspecifics and lions (Balme et al., 2017;Rafiq, 2016). ...
... Leopards sometimes consume kills on the ground but where intraguild competition for kills or carrion is high, they usually hoist their prey up trees to avoid being kleptoparasitized (Balme et al., 2017;Stein et al., 2015). Because lions do not balance well on the leading pole, we believe that the baits used in our study closely simulated feeding sites of leopards in the wild and that the observations closely reflect the species' natural behaviour at feeding sites. ...
... Intraspecific hostility can occur within and between sexes and among leopards of all ages (Farhadinia et al., 2018) and is associated with competition at kills (Steyn & Funston, 2006) or over territory (Balme & Hunter, 2004). Our findings of a strong competitive effect from male leopards were similar to a study by Balme et al. (2017) which showed that male leopards were responsible for 88% of intraspecific kleptoparasitism at Sabi Sand Game Reserve, South Africa. In the wild, antagonistic encounters between unfamiliar individuals occur (White & Harris, 1994) and the level of risk increases with increasing density, with associated competition for space, mates, and ...
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Knowledge of competition dynamics among Africa’s large carnivores is important for conservation. However, investigating carnivore behaviour in the field can be challenging especially for species that are difficult to access. Methods that enable remote collection of data provide a means of recording natural behaviour and are therefore useful for studying elusive species such as leopards (Panthera pardus). Camera traps and Global Positioning System (GPS) collars are powerful tools often used independently to study animal behaviour but where their data are combined, the interpretation of a species’ behaviours is improved. In this study we used data from baited camera trap stations to investigate the feeding habits of leopards at Malilangwe Wildlife Reserve, Zimbabwe. We investigated the influence of spotted hyenas, lions and other competing leopards on the feeding duration of leopards using Generalized Linear Mixed Effects Modelling. To test the influence of competing predators on resting distances from bait sites, eight leopards were fitted with GPS collars. Results showed that leopards spent the shortest time feeding on the baits in the presence of competing male leopards compared to other predators while lion presence caused animals to rest farthest from bait sites. Interaction analysis indicated that small‐bodied leopards spent significantly shorter durations feeding when spotted hyenas were present. Our findings demonstrate that competition from guild carnivores has negative impacts on the food intake of leopards, which may have implications for fitness and survival. This study provides a snapshot of the competition dynamics at bait sites which may give insight to ecosystem level interactions among large carnivores in savanna ecosystems.
... The results of our temporal spacing analysis suggest that leopard employ fine-scale avoidance of lion in areas where they co-occur in Ruaha-Rungwa. Lion are known to attack and kill leopard [20], steal kills from the smaller-bodied felid [67], and impact cub survival and recruitment rates in leopard populations [68]. However, multiple studies have found leopard to not spatially segregate from lion over larger areas [20,21,22]. ...
... In addition to the fine-scale temporal avoidance observed here, a number of additional coexistence mechanisms have been observed in other leopard populations, including fine-scale spatial avoidance in the presence of lion [9,19], avoiding areas where the probability of encountering lion is highest [20], minor temporal partitioning [21], dietary niche differentiation [70], and prey caching to avoid kleptoparasitism [67,71]. Thus, although this study provides evidence of fine-scale spatiotemporal avoidance and possible minor temporal partitioning, it is likely that leopard in Ruaha-Rungwa also make use of other adaptations that could not be identified by this study. ...
... This mirrors the results of a similar study in South Africa [58], and is likely to be a behaviour employed by spotted hyaena to increase kleptoparasitism and scavenging opportunities. Kleptoparasitism by spotted hyaena is often mainly targeted at leopard [9], and previous research has found spotted hyaena to be responsible for 50% of kleptoparasitism suffered by the species [67]. High rates of kleptoparasitism are associated with reduced reproductive success among female leopards [67]; the loss of prey to competitors, and particularly to spotted hyaena, therefore represents a threat to leopard fitness. ...
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Africa is home to some of the world’s most functionally diverse guilds of large carnivores. However, they are increasingly under threat from anthropogenic pressures that may exacerbate already intense intra-guild competition. Understanding the coexistence mechanisms employed by these species in human-impacted landscapes could help shed light on some of the more subtle ways in which humans may impact wildlife populations, and inform multi-species conservation planning. We used camera trap data from Tanzania’s Ruaha-Rungwa landscape to explore temporal and spatiotemporal associations between members of an intact East African large carnivore guild, and determine how these varied across gradients of anthropogenic impact and protection. All large carnivores except African wild dog ( Lycaon pictus ) exhibited predominantly nocturnal road-travel behaviour. Leopard ( Panthera pardus ) appeared to employ minor temporal avoidance of lion ( Panthera leo ) in all sites except those where human impacts were highest, suggesting that leopard may have been freed up from avoidance of lion in areas where the dominant competitor was less abundant, or that the need for leopard to avoid humans outweighed the need to avoid sympatric competitors. Lion appeared to modify their activity patterns to avoid humans in the most impacted areas. We also found evidence of avoidance and attraction among large carnivores: lion and spotted hyaena ( Crocuta crocuta ) followed leopard; leopard avoided lion; spotted hyaena followed lion; and lion avoided spotted hyaena. Our findings suggest that large carnivores in Ruaha-Rungwa employ fine-scale partitioning mechanisms to facilitate coexistence with both sympatric species and humans, and that growing human pressures may interfere with these behaviours.
... A 40-year dataset from the Sabi Sand Game Reserve, South Africa, revealed that average parental care to litters was 18 months, although this varied substantially from 9 to 35 months (Balme, Robinson, Pitman & Hunter, 2017). In addition, Balme et al. (2017) found that female leopards displayed extended parental care when prey were scarce, while providing prolonged care for sons. Dispersal may also be delayed by several months if food is abundant or if conspecifics occur at high densities . ...
... Leopards use camouflage and vegetation to stalk very close to their prey before sprinting, up to a speed of 60 km/h within 120 m and pouncing on their prey (Betram, 1979;. Hunting success varies from 5% in the Serengeti to 38% in Kaudom and captured prey are sometime hauled into trees out of reach of other large carnivores, or cached beneath vegetation Balme, Miller, Pitman & Hunter, 2017). Leopards have the most varied diet of any other African large carnivore, with a total of 92 prey species recorded in their diet in sub-Saharan Africa . ...
... For example, hunting bushbuck, common duiker, nyala and reedbuck also involves less risk and are generally easier to subdue . However, Balme et al. (2017) argue that leopards could select even smaller-than-expected prey species, as predicted by the optimal foraging theory, to balance trade-offs between losses (encountered through kleptoparasistism) and energetic gains (obtained by killing larger prey). ...
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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.
... Trinkel and Kastberger 2005;Watts and Holekamp 2008;M'soka et al. 2016;Sogbohossou et al. 2018) and lions and leopards (e.g. du Preez 2014; Stein et al. 2015;Balme et al. 2017a). Little research has been published on the effects of leopard activity on spotted hyaena activity (or vice versa), even though these two species have reported temporal activity overlaps of between 80% (Hayward and Slotow 2009) and 90% (Comley et al. 2020b). ...
... Lack of spatial avoidance could also be due to the cryptic colouration of leopards, allowing them to remain undetected for longer (Miller et al. 2018) or due to a low incurred cost of inhabiting similar areas (Balme et al. 2017b). Additionally, through behavioural changes such as prey caching, leopards can avoid interference competition by hoisting kills into trees (Stein et al. 2015;Balme et al. 2017a). ...
... Alternatively, patterns seen at the spatial and temporal level (spatial and temporal overlap instead of avoidance) could be driven by resource availability and behavioural plasticity (Maputla et al. 2015;Périquet et al. 2015;Rafiq et al. 2020). For example, by seeking cover and caching prey in trees (Balme et al. 2017a), leopards might not need to change their spatial and temporal patterns to avoid spotted hyaenas actively. ...
Article
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.
... 3. The abundance of alternative prey (e.g., non-ungulate species) inversely correlates with kill rate (i.e., prey switching for an abundant alternative prey reduces predation on primary prey; Elbroch et al. 2015a;Keehner et al. 2015;Soria-Díaz et al. 2018). 4. Puma kill rates positively correlate with species richness of scavengers that are dominant to pumas, as interspecific kleptoparasitism sometimes drives pumas to kill additional prey (Krofel et al. 2012;Elbroch and Wittmer 2013b;Elbroch et al. 2015b;Allen et al. 2021). 5. Kill rates of pumas might be higher at high puma densities, possibly to compensate for exploitative competition (e.g., if intraspecific kleptoparasitism is high, as documented for other felids at high density; Balme et al. 2017). Alternatively, high resources may attenuate intraspecific competition allowing carnivores to shrink home ranges and live at high densities (Šálek et al. 2015), or pumas mitigate competition by exhibiting complex social interactions (Elbroch et al. 2017b). ...
... Moreover, pumas are large carnivores that generally exist at low densities and intraspecific kleptoparasitism is probably infrequent in many populations due to the presumably low chances of encountering kills made by conspecifics. In contrast, high levels of carcass takeover by adult males, the dominant reproductive class, have been documented at high densities for another large felid, the leopard (Balme et al. 2017). However, recent data have challenged the view that pumas have few intraspecific interactions, and revealed that adult female pumas may commonly provide subsidies to resident adult males, which may be a potential investment in protection against non-resident males and infanticide (Elbroch et al. 2017b). ...
Article
Kill rates and functional responses are fundamental to the study of predator ecology and the understanding of predatory-prey dynamics. As the most widely distributed apex predator in the western hemisphere, pumas ( Puma concolor ) have been well studied, yet a synthesis of their kill rates is currently lacking. We reviewed the literature and compiled data on sex- and age-specific kill rate estimates of pumas on ungulates, and conducted analyses aimed at understanding ecological factors explaining the observed spatial variation. Kill rate studies on pumas, while numerous, were primarily conducted in Temperate Conifer Forests (< 10% of puma range), revealing a dearth of knowledge across much of their range, especially from tropical and subtropical habitats. Across studies, kill rates in ungulates/week were highest for adult females with kitten(s) (1.24 ± 0.41 ungulates/week) but did not vary significantly between adult males (0.84 ± 0.18) and solitary adult females (0.99 ± 0.26). Kill rates in kg/day differed only marginally among reproductive classes. Kill rates of adult pumas increased with ungulate density, particularly for males. Ungulate species richness had a weak negative association with adult male kill rates. Neither scavenger richness, puma density, the proportion of non-ungulate prey in the diet, nor regional human population density had a significant effect on ungulate kill rates, but additional studies and standardization would provide further insights. Our results had a strong temperate-ecosystem bias highlighting the need for further research across the diverse biomes pumas occupy to fully interpret kill rates for the species. Data from more populations would also allow for multivariate analyses providing deeper inference into the ecological and behavioural factors driving kill rates and functional responses of pumas, and apex predators in general.
... Contrary to previous studies in Thailand that showed the diet of Indochinese leopard was dominated by relatively small prey, including primates, small ungulates and small carnivores [16][17][18], the leopard diet in SWS was dominated by large (greater than 30 kg) ungulates (64.3% BC), which did not support our prediction. In Africa, leopard also feed mainly on large ungulates in open habitats [59,[74][75][76][77][78][79]. Leopard may have had similar dietary niche in EPL as the African sites because open habitats typically have a higher carrying capacity for large ungulates than closed forests, resulting in leopard predating more on large ungulates in open habitats due to their higher availability and/or accessibility. ...
... Our results regarding large differences in male and female diets were unexpected, given that previous studies in Africa did not find major differences in body size of ungulates consumed by male and female leopard [59,61]. In one exception, Balme et al. [74] found male leopard consumed prey that was on average one-third larger than prey consumed by females. By contrast, our results showed that the main prey consumed by male leopard in SWS was probably 12-30 times larger (depending on age of prey) than the main prey consumed by females. ...
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We studied the Indochinese leopard (Panthera pardus delacouri) in eastern Cambodia, in one of the few potentially remaining viable populations in Southeast Asia. The aims were to determine the: (i) current leopard density in Srepok Wildlife Sanctuary (SWS) and (ii) diet, prey selection and predation impact of leopard in SWS. The density, estimated using spatially explicit capture–recapture models, was 1.0 leopard/100 km², 72% lower than an estimate from 2009 at the same site, and one of the lowest densities ever reported in Asia. Dietary analysis of 73 DNA confirmed scats showed leopard consumed 13 prey species, although ungulates comprised 87% of the biomass consumed (BC). The overall main prey (42% BC) was banteng (Bos javanicus), making this the only known leopard population whose main prey had adult weight greater than 500 kg. Consumption of wild pig (Sus scrofa) was also one of the highest ever reported (22% BC), indicating leopard consistently predated on ungulates with some of the largest adult weights in SWS. There were important differences in diet and prey selection between sexes, as males consumed mostly banteng (62% BC) in proportion to availability, but few muntjac (Muntiacus vaginalis; 7% BC), whereas females selectively consumed muntjac (56% BC) and avoided banteng (less than 1% BC). Predation impact was low (0.5–3.2% of populations) for the three ungulate species consumed. We conclude that the Indochinese leopard is an important apex predator in SWS, but this unique population is declining at an alarming rate and will soon be eradicated unless effective protection is provided.
... Ultimately, local displacement (Newsome et al., 2017) and a reduction in resource availability due to exploitative competition (Caro & Stoner, 2003) influence carnivore HR size. Local-scale studies have shown inconsistent results regarding the role of dominant carnivores on leopard population density and activity (Steinmetz et al., 2013;Vanak et al., 2013;Balme et al., 2017). However, no attempt has been made so far to understand general trends on how the presence of different competitors influences the HR behaviour of leopards on a global scale. ...
... Interspecific interactions are major drivers shaping the structure of ecological communities (Hardin, 1960;Krebs, 2014), and local-scale studies have revealed a variable influence of intraguild competition on leopard abundance (Steinmetz et al., 2013;Vanak et al., 2013;Balme et al., 2017). At a global scale, our models showed a non-significant effect of competitors on HR size. ...
Article
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Movement is a fundamental process in animal ecology. For many species, such as large carnivores, movement patterns are greatly shaped by a combination of ecological and anthropogenic factors. Understanding how these factors impact the roaming capacity of large carnivores is essential to forecast risks and design long‐term conservation strategies. The leopard (Panthera pardus) is a generalist predator broadly distributed over varied and different environments, but global leopard populations are declining at a concerning rate and conservation actions are pressing. This scenario makes the leopard a suitable species to understand how global ecological and anthropogenic drivers affect the spatial behaviour of large carnivores and how these should inform conservation efforts. We compiled data from local studies worldwide and used macroecological (climatic, productivity, and human footprint), and intra‐ and interspecific (conspecifics, competitors and prey) predictors to model the roaming requirements of leopards based on home range sizes. Male home range size was largely and positively related to the range sizes of local females and inversely to vegetation productivity. For females, higher seasonal variations in temperature like the observed in arid areas were associated with larger home ranges, while increased human impact resulted in smaller home ranges likely due to concentrated food resources such as domestic species. These predictors are linked to threatening global change processes due to anthropogenic activities that will likely impact the roaming behaviour of leopards in the coming decades with potential consequences for their populations worldwide. Our results provide crucial information towards the development of integrative research linking macroecological and local variables with global change predictions that can inform conservation programmes addressing future risks of degradation, endangerment and human‐leopard conservation conflicts. The leopard (Panthera pardus) is an iconic, generalist and broadly distributed felid undergoing severe population declines. We conducted macroecological research to identify the determinants of leopard home range size across their global distribution. Increasing climate warming and landscape anthropization will strongly impact leopard movements and home range size, which is highly significant for transboundary conservation efforts.
... Additionally, species that demonstrate preferred foraging times (e.g., the leopard seal; Krause et al. 2016) may decouple prey acquisition and consumption to maximize access during peak periods (Sherry 1985). This strategy may also reduce kleptoparasitism, or direct prey theft, especially for short-term hoarders (Balme et al. 2017). Therefore, temporal decoupling is a likely driver of caching behavior in high density leopard seal foraging grounds where kleptoparasitism has been consistently observed (Hiruki et al. 1999; Vera ...
... As intraspecific competition increases and larder hoarding becomes untenable, the prevalence of hiding prey underwater and away from prey acquisition sites is likely to increase (Vander Wall 1990). Finally, we expect caching to be more prevalent in smaller, less dominant animals that hoard to gain competitive advantage and reduce the incidence of kleptoparasitism (Balme et al. 2017). Overall, it is likely that variability in the leopard seal's polar marine habitat and conspecific density necessitates a greater diversity of caching strategies compared with terrestrial bioregions. ...
Article
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The foraging behaviors of apex predators can fundamentally alter ecosystems through cascading predator–prey interactions. Food caching is a widely studied, taxonomically diverse behavior that can modify competitive relationships and affect population viability. We address predictions that food caching would not be observed in the marine environment by summarizing recent caching reports from two marine mammal and one marine reptile species. We also provide multiple caching observations from disparate locations for a fourth marine predator, the leopard seal (Hydrurga leptonyx (de Blainville, 1820)). Drawing from consistent patterns in the terrestrial literature, we suggest the unusual diversity of caching strategies observed in leopard seals is due to high variability in their polar marine habitat. We hypothesize that caching is present across the spectrum of leopard seal social dominance; however, prevalence is likely to increase in smaller, less-dominant animals that hoard to gain competitive advantage. Given the importance of this behavior, we draw attention to the high probability of observing food caching behavior in other marine species.
... The outcome of interspecific interactions also depends on the relative rank of the predators within the "intraguild dominance hierarchy" (Groom et al., 2017), which is driven in part by competitors relative body size and social structure. In the African savannah, wild dog (Lycaon pictus), cheetah (Acinonyx jubatus), and leopard (Panthera pardus) prematurely abandon their kills when larger-bodied, social predators, such as lion and/or hyena, are present (Balme et al., 2017;Fanshawe & Fitzgibbon, 1993;Hunter et al., 2007). In northern systems, bear presence causes mountain lions (Puma concolor) and Eurasian lynx (Lynx lynx) to abandon their kills (Allen et al., 2021;Elbroch et al., 2014;Engebretsen et al., 2021;Krofel et al., 2012), and wolves similarly displace mountain lions from kill sites (Ruth et al., 2019). ...
Article
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Competition between apex predators can alter the strength of top‐down forcing, yet we know little about the behavioral mechanisms that drive competition in multipredator ecosystems. Interactions between predators can be synergistic (facilitative) or antagonistic (inhibitive), both of which are widespread in nature, vary in strength between species and across space and time, and affect predation patterns and predator‐prey dynamics. Recent research suggests gray wolf (Canis lupus) kill rates decrease where they are sympatric with brown bears (Ursus arctos), however, the mechanisms behind this pattern remain unknown. We used data from two long‐term research projects in Scandinavia (Europe) and Yellowstone National Park (North America) to test the role of interference and exploitation competition from bears on wolf predatory behavior, where altered wolf handling and search time of prey in the presence of bears are indicative of interference and exploitation competition, respectively. Our results suggest the mechanisms driving competition between bears and wolves were dependent on the season and study system. During spring in Scandinavia, interference competition was the primary mechanism driving decreased kill rates for wolves sympatric with bears; handling time increased, but search time did not. In summer, however, when both bear and wolf predation focused on neonate moose, the behavioral mechanism switched to exploitation competition; search time increased, but handling time did not. Interference competition, however, did affect wolf predation dynamics in Yellowstone during summer, where wolves prey more evenly on neonate and adult ungulates. Here, bear presence at a carcass increased the amount of time wolves spent at carcasses of all sizes and wolf handling time for small prey, but decreased handling time for the largest prey. Wolves facilitate scavenging opportunities for bears, however, bears alter wolf predatory behavior via multiple pathways and are primarily antagonistic to wolves. Our study helps clarify the behavioral mechanisms driving competition between apex predators, illustrating how interspecific interactions can manifest into population‐level predation patterns.
... This supports the suggestion that leopards are largely unaffected by competition for food resources with other large predators (Hayward and Kerley, 2008). Leopards have been shown to be able to cope with kleptoparasitism by spotted hyaenas through temporal avoidance (Havmøller et al., 2020) and by caching kills out of reach in trees (Balme et al., 2017). Coupled with this, leopard body size overlaps sufficiently with that of hyaenas to limit interference competition via direct lethal confrontation (Bailey, 1993). ...
Article
Globally three quarters of large terrestrial mammalian predators are in decline and many populations are data deficient, including those of African leopards across much of their range. Here we assess the drivers of decline African leopard populations in 16 camera trap surveys covering a total area of 15,120 km², across a gradient of anthropogenic impact, management and geography, in protected areas across the Zimbabwean component of the Kavango-Zambezi (KAZA) Transfrontier Conservation Area. Population density was calculated using spatially explicit mark-recapture estimators and Generalised Additive Models (GAM) were used to assess factors affecting population density. Density estimates ranged from 0.7 to 12.2 (mean 2.9 ± 2.7) leopards/100km². Leopard density was higher in wooded sites and rugged terrain but negatively affected by human factors including human appropriation of net primary productivity (HANPP), trophy hunting risk and bush-meat poaching. High lion densities (>6.0 lions/ 100km²) negatively affected leopard density. Annual rainfall over a gradient of ~300 mm across survey sites was not influential in predicting population density. Previous assessments of the drivers of declining leopard population density (CITES 1988), asserting that leopard densities can be predicted by annual rainfall and are unaffected by human disturbance in unmodified habitat are not supported by our findings. We recommend that the 1988 assessment, used to manage CITES leopard trophy hunting export quotas since the late 1980s, should be reviewed.
... Spotted hyaenas were also absent from our study, due to no individuals being GPS collared over the study period. However, spotted hyaenas can have significant impacts on leopard fitness through kleptoparasitism (Balme et al., 2017a) and spatial capture-recapture studies suggest hyaenas actively track leopards (Balme et al., 2019). Further work on the factors predisposing leopard-hyaena encounters is also thus warranted. ...
Article
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. Encounters between individuals in the wild can have implications for a range of processes; yet for large carnivores, difficulties in collecting data mean relatively few studies have explored the factors contributing to direct encounters between species. Using high‐resolution GPS collar data, we explore the spatial and temporal factors impacting encounter occurrences between leopards and other large African carnivore species. Our results show how habitat characteristics and niche overlaps contribute to encounters between competitors and suggest how niche shifts within competitor assemblages could impact competition between species.
... Scavenging is an important ecosystem process (Wilson & Wolkovich 2011) and scavengers can importantly affect predation, reproduction, social system and evolution of predators (Cooper 1991, Iyengar 2008, Balme et al. 2017, Tallian et al. 2017, including Eurasian lynx (Mattisson et al. 2011, Krofel & Jerina 2016 (Krofel 2012), which in combination with GPS telemetry showed that lynx lost 32% of their kills and 15% of all consumable biomass to scavenging bears, which were managed to displace lynx from their kill sites and reduced the number of days lynx are able to feed . These values correspond very well to data obtained in this study (33% of lynx kill sites were found by bears; Table 1). ...
Article
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Recent technological developments in non-invasive methods enable efficient ways to study behaviour of elusive predators. We used two types of automatic digital video surveillance systems in combination with GPS telemetry to record feeding behaviour of wild Eurasian lynx (Lynx lynx), intraspecific prey sharing and scavenger activity at ungulate kill sites in Dinaric Mountains. This approach proved as very effective and mostly non-invasive way to obtain detailed data about the consumption of prey by lynx and kleptoparasites, especially when the advanced video system was used. Lynx spent considerable amount of time in the vicinity of the kill site, but usually visited the carcass for feeding only once per night with a mean visit time of 35 min and most of the feeding occurred during the first half of the night. Lynx covered 83% of prey remains, which seemed to be effective against avian scavengers that only found 17% of the carcasses. We recorded six vertebrate species scavenging on lynx kills, with red fox (Vulpes vulpes) and brown bear (Ursus arctos) being the most frequent kleptoparasites. Based on our experiences, we provide recommendations for future research using this method, as well as pros and cons of the advanced vs. simple video systems.
... During times of high prey density, multiple top predator species might have used areas of highly aggregated prey and thus increased encounter rates F I G U R E 2 Monthly survival for (a) adult and (b) cub cheetahs in relation to prey density and lion density during dry seasons of years with average resighting rates, Mun-Ya-Wana Conservancy, KwaZulu-Natal, South Africa, 2008 with cheetahs. In addition, lions occasionally kill and compete with leopards and hyaenas (Balme, Miller, Pitman, & Hunter, 2017;Trinkel & Kastberger, 2005), and leopards, similar to cheetahs, have been found to use fine-scale spatial partitioning to avoid interactions with lions (du Preez, Hart, Loveridge, & Macdonald, 2015;Vanak et al., 2013). ...
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1.Research on drivers of demographic rates has mostly focused on top predators and their prey, and comparatively less research has considered the drivers of mesopredator demography. Of those limited studies, most focused on top‐down effects of apex predators on mesopredator population dynamics, whereas studies investigating alternative mechanisms are less common. 2.In this study, we tested hypotheses related to top‐down, bottom‐up, and density‐dependent regulation of demographic rates in an imperiled mesopredator, the cheetah (Acinonyx jubatus). 3.We used a 25‐year dataset of lion density, cheetah density, and prey density from the Mun‐Ya‐Wana Conservancy in South Africa and assessed the effects of top‐down, bottom‐up, and density‐dependent drivers on cheetah survival and reproduction. 4.In contrast to the top‐down and bottom‐up predications, both adult and juvenile cheetahs experienced the lowest survival during months with high prey densities and low lion densities. We only observed support for a density‐dependent response in juvenile cheetahs, where they had a higher probability of reaching independence during times with low cheetah density, and low prey density. We did not identify any strong drivers of litter size. 5.Collectively, our results indicate that high apex predator abundance might not always have negative effects on mesopredator populations, and suggest that context dependency in top‐down, bottom‐up, and density‐dependent factors may regulate demographic rates of cheetahs and other mesopredators. Our results highlight the complexities of population‐level drivers of cheetah demographic rates and the importance of considering multiple hypotheses of mesopredator population regulation. This article is protected by copyright. All rights reserved.
... Besides, as suggested by Smith et al. (2015), an increase in ungulate carcasses abandoned by predators in disturbed areas may impact the entire biotic community, providing additional subsidies to scavengers. Finally, a reduced feeding time may also negatively influence reproductive success, as observed in leopards (Panthera pardus) even in areas with high prey densities (Balme et al., 2017). ...
Article
Over the past decades, non-consumptive outdoor recreation has intensified, resulting in a more widespread and regular human presence in natural habitats, including protected areas. This has shown to negatively affect several animal species, and in some cases, cause their decline. Therefore, understanding the impacts of recreation on protected species is fundamental. In the Bohemian Forest Ecosystem, we GPS-monitored the Eurasian lynx (Lynx lynx), generally considered tolerant to human presence. We tested whether the local level of recreation influenced (a) time spent by lynx at killed prey, both in terms of number of hours each night and of number of nights at each killed prey (i.e. feeding behavior) and (b) selection of daytime resting sites. Furthermore, we checked whether each behavior was influenced by local habitat features ensuring low accessibility to people and high protective cover, and by the level of nature protection assigned to different parts of the study area, all of which likely influence perceived risk by lynx. Finally, we tested for seasonal (winter vs. summer) changes in these variables' effects. Throughout the year, the local intensity and recurrence of recreation was negatively correlated with the probability that lynx would use a given location for daytime resting and with the number of hours that lynx spent at a given killed prey each night. Furthermore, habitat features providing protective cover positively correlated with both behaviors, and the probability that lynx would use a given location for daytime resting was higher inside than outside protected areas. Finally, recreation negatively correlated with the number of nights lynx spent at killed prey only in winter (i.e. October-April). These findings can be applied when planning recreational activities, and generally highlight the need for a deeper understanding of the impacts of human activities across a range of species.
... But cooperative feeding often comes with a cost; food captured must be shared, limiting the potential resource gain for each individual involved (Sachs et al. 2004). Hunting close to conspecifics, whether cooperatively or not, also provides opportunity for food stealing or kleptoparasitism, meaning that predators may need to defend their kills, or relocate and hide prey to prevent others taking it (Balme et al. 2017). In this case, the predator must incur the extra cost of these behaviours in addition to the original hunting and eating costs (Schoener 1971;Pyke et al. 1977). ...
Article
Cooperative feeding is often observed among predators with strong social bonds; however, it is unexpected in solitary predators. During 2016, several mass predation events were witnessed in St Andrews Bay and Right Whale Bay, South Georgia, where up to 36 leopard seals (Hydrurga leptonyx) were seen feeding together at king penguin (Aptenodytes patagonicus) colonies. Three post-mortem prey-processing events were observed where two leopard seals actively fed on the same carcass in an unusual display of tolerance for a species where anti-social behaviour is the norm. The seals were observed repeatedly tearing adult king penguins between themselves, while floating alongside each other at the surface of the water. This is the first record of co-feeding in this difficult-to-study species; however, it is expected that the behaviour is rare within the population. We propose that the high density of predators combined with the readily available prey, makes it costlier to defend a kill than it is to tolerate kleptoparasitism. It is unclear whether this behaviour shows cooperative feeding, which would likely enable more efficient prey processing: by holding the prey in their jaws, each seal provides an anchor on the prey that others can pull against to stretch and tear it.
... Cabe destacar la conducta de almacenamiento de las presas (L. europaeus) por el gato de Geoffroy en la comuna de Coyhaique durante el invierno, ya que ésta no había sido reportada anteriormente y podría relacionarse con la satisfacción de sus demandas energéticas durante la temporada invernal (Sandell, 1989) o para evitar el cleptoparasitismo (Balme, Miller, Pitman, Hunter, 2017). ...
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Reportamos cinco registros directos del gato de Geoffroy Leopardus geoffroyi, en las zonas sur y austral de Chile. Los registros se presentaron en diferentes hábitats, tanto en sitios con vegetación nativa y con presencia humana, en donde se presentan amenazas para su supervivencia. Adicionalmente, se consultó la literatura disponible publicada y bases de datos para corroborar los registros previos de esta especie en Chile. Los registros aquí reportados contribuyen a fortalecer la información sobre la distribución del gato de Geoffroy en Chile, sin embargo, es indispensable realizar más investigaciones con el fin de conocer su situación poblacional actual, amenazas y soluciones.
... This has been observed in other carnivores, such as leopards (P. pardus), where females that suffered from a high rate of kleptoparasitism had lower reproductive success (Balme, Miller, Pitman, & Hunter, 2017). However, the chance of falling victim to kleptoparasitism is reduced in areas with dense vegetation (Hunter, Durant, & Caro, 2007a;Paulson, 1985) and it is possible that areas with open habitat experience reduced cub recruitment due to predation and kleptoparastism. ...
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Recruitment is a critical parameter governing population dynamics and influences population persistence. Understanding the drivers of recruitment is therefore important for conservation, especially for long-lived mammals such as large carnivores, which have low reproductive rates, rendering them prone to extinction. Using cheetahs (Acinonyx jubatus) as a model species, I investigated the variation in cub recruitment in relation to habitat and the abundance of tourists and predators. Per litter, female cheetahs on average raised 1.71 ± 1.35 cubs to independence, but this varied depending on the presence of open habitat and the abundance of tourists, both of which had a negative effect on cub recruitment. More specifically, female cheetahs that were mostly found in open habitats on average raised 1.69 ± 0.14 cubs per litter to independence compared to 3.04 ± 0.26 cubs in denser habitat. Similarly, female cheetahs that were exposed to high tourist abundance on average raised 0.21 ± 0.72 cubs to independence compared to 2.32 ± 0.11 cubs in low tourism areas. Neither lion nor spotted hyaena abundance had an impact on the number of cubs that were recruited. Based on these findings, I recommend that the importance of a heterogeneous environment should be taken into consideration in habitat management, restoration efforts, and reintroduction programs. In addition, tourist quotas should be put in place in high visitation areas and strict wildlife viewing guidelines, such as number of vehicles, tourist behavior, time spent, and distance to a sighting, should be enforced. Cub recruitment is an important component of species persistence and incorporating these findings could aid conservation efforts for species that are increasingly under threat.
... When hunting large prey, maximizing nutritional gain requires spending substantial time handling the carcass, which increases the time spent in a risky situation. Moving the kill to a refuge is a strategy used by leopards (Panthera pardus) to lower rates of kleptoparasitism (Balme et al. 2017), while pumas (Puma concolor) cache large carcasses making their kills less likely to be detected by bears (Ursus americanus and arctos) (Murphy et al. 1998). Cheetahs cannot conceal their prey nor can they reliably defend their kills against larger predators and therefore they must employ different strategies, wile lions and hyenas are more likely to find and steal larger kills (Hunter et al. 2007b). ...
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While handling large kills, mesocarnivores are particularly vulnerable to kleptoparasitism and predation from larger predators. We used 35 years of observational data on cheetah (Acinonyx jubatus) hunts in Serengeti National Park to investigate whether cheetahs’ prey handling behavior varied in response to threats from lions (Panthera leo) and spotted hyenas (Crocuta crocuta). Male cheetahs and single females, whose main threat was kleptoparasitism, minimized time on the kill by being less vigilant and eating quickly, thereby shortening their handling times. Mothers with cubs showed a different strategy that prioritized vigilance over speed of eating, which increased time spent handling prey. Vigilance allowed them to minimize the risk of their cubs being killed while giving cubs the time they need to eat at the carcass. Flexible behavioral strategies that minimize individual risk while handling prey likely allow mesocarnivores to coexist with numerous and widespread apex predators. Significance statement Medium-sized carnivores like cheetahs face the challenge of coexisting with larger carnivores that steal their kills and kill their cubs. We investigated how cheetahs modify their behavior on kills to minimize risks from larger predators. Using 35 years of data on 400+ cheetah hunts across 159 individuals, we found that cheetahs without cubs whose primary danger is having their kill stolen spent little time engaged in vigilance and instead ate quickly, reducing the risk of theft. Mothers with cubs, however, took a slower approach and were more vigilant while handling prey to avoid cub predation by lions and spotted hyenas. The ability of cheetahs to modify their prey handling behavior depending on the type of risk they face likely allows them to coexist with numerous larger carnivores.
... Research directly linking bear kleptoparatism to puma fitness, however, is lacking. For example, kleptoparatism of leopard (Panthera pardus) kills negatively impacts leopard reproductive success (e.g., Balme et al., 2017). At this time, we cannot currently predict whether there exists an energetic threshold at which point bear kleptoparatism impacts puma fitness, or say definitively that it does. ...
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Background Interspecific competition affects species fitness, community assemblages and structure, and the geographic distributions of species. Established dominance hierarchies among species mitigate the need for fighting and contribute to the realized niche for subordinate species. This is especially important for apex predators, many of which simultaneous contend with the costs of competition with more dominant species and the costs associated with human hunting and lethal management. Methods Pumas are a widespread solitary felid heavily regulated through hunting to reduce conflicts with livestock and people. Across their range, pumas overlap with six apex predators (gray wolf, grizzly bear, American black bear, jaguar, coyote, maned wolf), two of which (gray wolf, grizzly bear) are currently expanding in North America following recovery efforts. We conducted a literature search to assess whether pumas were subordinate or dominant with sympatric apex predators, as well as with three felid mesocarnivores with similar ecology (ocelot, bobcat, Canada lynx). We also conducted an analysis of the spatial distributions of pumas and their dominant sympatric competitors to estimate in what part of their range, pumas are dominant versus subordinate. Results We used 64 sources to assess dominance among pumas and other apex predators, and 13 sources to assess their relationships with felid mesocarnivores. Evidence suggested that wolves, grizzly bears, black bears, and jaguars are dominant over pumas, but that pumas are dominant over coyotes and maned wolves. Evidence suggested that pumas are also dominant over all three felid mesocarnivores with which they share range. More broadly, pumas are subordinate to at least one other apex carnivore in 10,799,252 (47.5%) of their 22,735,268 km ² range across North and South America. Discussion Subordinate pumas change their habitat use, suffer displacement at food sources, likely experience increased energetic demands from harassment, exhibit increased starvation, and are sometimes directly killed in competitive interactions with dominant competitors. Nevertheless, we lack research clearly linking the costs of competition to puma fitness. Further, we lack research that assesses the influence of human effects simultaneous with the negative effects of competition with other sympatric carnivores. Until the time that we understand whether competitive effects are additive with human management, or even potentially synergistic, we encourage caution among managers responsible for determining harvest limits for pumas and other subordinate, apex carnivores in areas where they are sympatric with dominant species. This may be especially important information for managers working in regions where wolves and brown bears are recolonizing and recovering, and historic competition scenarios among multiple apex predators are being realized.
... It is noteworthy that areas of high density between the species do not appear to overlap (Fig. 3). Previous studies have suggested that spotted hyenas can be significant kleptoparasites of leopard kills, forcing them to cache or avoid areas with high hyena density (Balme et al., 2017a;Davis et al., 2021). Similarly, another study detected low temporal overlap between leopards and spotted hyenas in Tanzania, which was postulated to be due to the avoidance of kleptoparasitism (Havmøller et al., 2020). ...
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Robust measures of animal densities are necessary for effective wildlife management. Leopards (Panthera pardus) and spotted hyenas (Crocuta Crocuta) are higher order predators that are data deficient across much of their East African range and in Uganda, excepting for one peer-reviewed study on hyenas, there are presently no credible population estimates for these species. A lack of information on the population status and even baseline densities of these species has ramifications as leopards are drawcards for the photo-tourism industry, and along with hyenas are often responsible for livestock depredations from pastoralist communities. Leopards are also sometimes hunted for sport. Establishing baseline density estimates for these species is urgently needed not only for population monitoring purposes, but in the design of sustainable management offtakes, and in assessing certain conservation interventions like financial compensation for livestock depredation. Accordingly, we ran a single-season survey of these carnivores in the Lake Mburo National Park of southwestern Uganda using 60 remote camera traps distributed in a paired format at 30 locations. We analysed hyena and leopard detections under a Bayesian spatially explicit capture-recapture (SECR) modelling framework to estimate their densities. This small national park (370 km 2) is surrounded by Bahima pastoralist communities with high densities of cattle on the park edge (with regular park incursions). Leopard densities were estimated at 6.31 individuals/100 km 2 (posterior SD = 1.47, 95% CI [3.75-9.20]), and spotted hyena densities were 10.99 Distributed under Creative Commons CC-BY 4.0 individuals/100 km 2 , but with wide confidence intervals (posterior SD = 3.35, 95% CI [5.63-17.37]). Leopard and spotted hyena abundance within the boundaries of the national park were 24.87 (posterior SD 7.78) and 39.07 individuals (posterior = SD 13.51) respectively. Leopard densities were on the middle end of SECR studies published in the peer-reviewed literature over the last 5 years while spotted hyena densities were some of the first reported in the literature using SECR, and similar to a study in Botswana which reported 11.80 spotted hyenas/100 km 2. Densities were not noticeably lower at the park edge, and in the southwest of our study site, despite repeated cattle incursions into these areas. We postulate that the relatively high densities of both species in the region could be owed to impala Aepyceros melampus densities ranging from 16.6-25.6 impala/km 2. Another, potential explanatory variable (albeit a speculative one) is the absence of interspecific competition from African lions (Panthera leo), which became functionally extinct (there is only one male lion present) in the park nearly two decades ago. This study provides the first robust population estimate of these species anywhere in Uganda and suggests leopards and spotted hyenas continue to persist in the highly modified landscape of Lake Mburo National Park.
... While many large carnivores provide leftovers to scavengers, some large carnivores are more takers than givers, stealing kills from smaller carnivores rather than (or in addition to) providing them. For some mid-ranked carnivores such as cheetahs and leopards, avoidance of kleptoparisitism is a driving force in their ecology (Durant 2000b;Scantlebury et al. 2014;Balme et al. 2017;Hilborn et al. 2018). Kleptoparasitism has primarily been examined within east African carnivore communities; 7 of the 12 studies in our analysis occurred in Africa. ...
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Interactions among terrestrial carnivores involve a complex interplay of competition, predation and facilitation via carrion provisioning, and these negative and positive pathways may be closely linked. Here, we developed an integrative framework and synthesized data from 256 studies of intraguild predation, scavenging, kleptoparisitism and resource availability to examine global patterns of suppression and facilitation. Large carnivores were responsible for one third of mesocarnivore mortality (n = 1,581 individuals), and intraguild mortality rates were superadditive, increasing from 10.6% to 25.5% in systems with two vs. three large carnivores. Scavenged ungulates comprised 30% of mesocarnivore diets, with larger mesocarnivores relying most heavily on carrion. Large carnivores provided 1,351 kg of carrion per individual per year to scavengers, and this subsidy decreased at higher latitudes. However, reliance on carrion by mesocarnivores remained high, and abundance correlations among sympatric carnivores were more negative in these stressful, high-latitude systems. Carrion provisioning by large carnivores may therefore enhance suppression rather than benefiting mesocarnivores. These findings highlight the synergistic effects of scavenging and predation risk in structuring carnivore communities, suggesting that the ecosystem service of mesocarnivore suppression provided by large carnivores is strong and not easily replaced by humans.
... One of the potential interactions between jackals and local wildlife that remains unstudied is kleptoparasitism, also known as food stealing (Iyengar, 2008). This includes scavenging on prey killed by apex predators, which can cause considerable food loss for the predators (Allen et al., 2015;Krofel and Jerina, 2016), indirectly increase kill rates (Krofel et al., 2012), reduce reproductive success (Balme et al., 2017), or even pose a threat for the predator population (Gorman et al., 1998). It has been also suggested that, in the long term, kleptoparasitism may trigger changes in prey choice, social system, and evolution of predators (Iyengar, 2008;Krofel et al., 2012). ...
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The arrival of a new carnivore can have important effects on local communities. While several effects of introduced alien species have been well documented, few studies have reported the ecological consequences of an expanding native species. Golden jackals (Canis aureus) are rapidly expanding their distribution in Europe, far beyond their historic range. While this raises many concerns about their potential impact on native wildlife, actual consequences are rarely recorded. Besides being a predator, the jackal is also an efficient scavenger and could function as a kleptoparasite for other predators living in areas colonized by jackals. Large felids are among the predators most vulnerable to kleptoparasitism and Eurasian lynx (Lynx lynx) are already known to be negatively affected by several scavengers. Here we report on the first confirmed cases of jackals scavenging on lynx kills in the Dinaric Mountains, Slovenia. We used camera traps to monitor scavengers at 65 lynx kills and recorded two cases of groups of jackals feeding on roe deer killed by lynx. To determine the potential for jackal kleptoparasitism on lynx at the continental level, we also calculated trends in the overlap in distribution ranges of both species in Europe. To date, jackals have colonized 13% of lynx range, including parts of two highly threatened populations. Finally, we highlight the potential impact of sympatric grey wolves (Canis lupus) to modulate this newly described jackal-lynx kleptoparasitic interaction.
... since fresh carcasses often attract several scavengers, which results in increased probability for agonistic inter-or intra-specific interactions (Allen, Wilmers, Elbroch, Golla, & Wittmer, 2016). Many scavengers and predators have developed various behavioral adjustments to reduce the competition, such as caching the carcass in trees, in caves or covering it with various materials (Balme, Miller, Pitman, & Hunter, 2017;Bischoff-Mattson & Mattson, 2009;de Ruiter & Berger, 2001;Krofel, Skrbinšek, & Mohorovi c, 2019). ...
Article
While scavenging has been repeatedly reported for several felid species, surprisingly little information is available on scavenging behavior of the European wildcat (Felis silvestris). To fill this knowledge gap, we used camera traps to document scavenging behavior at the 48 experimentally-set deer carcasses at random locations throughout the year. We recorded European wildcats scavenging on 38% of the eight carcasses set in winter and on none set in the other parts of the year. Wildcats fed on two carcasses for extended periods (up to 22 days) with an average of 3.3 visits per day and 7.8-h interval between the visits. We recorded scavenging throughout the day, but analysis indicated a crepuscular pattern. We also recorded caching behavior on 7% of the visits (n = 105), when wildcats used leaves or snow to partly or completely cover the carcasses. Beside wildcats, 12 other vertebrate species of scavengers were recorded at the carcasses. We recorded agonistic interaction with European badger (Meles meles) and despite its smaller size, the wildcat managed to defend the carcass. The extensive feeding, frequent caching behavior and active defense from scavengers indicate that the wildcats recognized the ungulate carcasses as an important food source in winter and that scavenging could be a neglected aspect of the European wildcat ecology. We also suggest that caching behavior could be regularly used by the European wildcat when feeding on larger carcasses, but was likely previously missed due to limited research effort to record scavenging and caching behavior.
... Felids display caching behaviour in multiple ways. Although leopards (Panthera pardus) can 41 cache their prey into caves (Ruiter and Berger 2001), they more commonly carry prey up to 42 trees to reduce the risk of kleptoparasitism from not-climbing competitors (Balme et al. 2017). 43 ...
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Caching behavior consists on the relocation or storage of food to protect it from competitors, to delay food spoilage, or to exploit it during times of scarcity. While this behavior has been widely described for some medium and large-sized felids, only a few cases documented caching behavior in small felids. Here, we provide the first exhaustive description of a caching event on a European wildcat in the Cantabrian Mountains (NW Spain). The wildcat behaved like a lynx/puma, visiting a road-killed roe deer carcass at least 9 days along a 21-day period, consuming the main muscles and covering it with hair and vegetation.
... Caching has been observed in many taxonomic groups; in mammals, well-known examples include many species of rodents caching nuts or seeds (Smith and Reichman 1984) and Arctic foxes (Vulpes lagopus) caching excess eggs for later consumption during a limited waterfowl breeding season (Careau et al. 2007). Many large carnivores cache the remainder of ungulate or other large kills that are too large to consume in a single feeding bout, either in a tree (e.g., leopards [Panthera pardus]; Balme et al. 2017) or covered with soil or other debris for later consumption (e.g., pumas [Puma concolor]; Hornocker 1970). Although caching of food has been observed in many carnivores, there is variation among species in how and why caching is used. ...
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Food caching is a common behavior for many mammals, but less is known about the prevalence and importance of food caching for some species. Here we report the first documented caching events by Asiatic black bears (Ursus thibetanus, n = 5) and three additional caching events by American black bears (U. americanus). We also performed a systematic literature review on caching by bears as a reference point for future investigations. Caching was most frequently reported for brown bears (U. arctos), and most caching by bears occurred with large prey. Caching is most likely used to protect large carcasses from spoiling or detection by scavengers, allowing bears to consume more of the carcass. The lack of published studies on caching by bears may be due to the behavior being infrequently used and difficult to record. We encourage an increase, but also consistency, in future reporting, including specific descriptions of caching behavior.
... bury or partially bury (i.e. short-term caching) their prey items after an initial feeding bout (Balme, Miller, Pitman, & Hunter, 2017). One explanation for this behaviour is the food-perishability hypothesis, which postulates that short-term cachers store food to deter or delay food spoilage, that is to manipulate microbial growth (Bischoff-Mattson & Mattson, 2009). ...
Article
1. Carrion is long recognized as important to scavengers. How carrion may affect soil microbial biodiversity and ecosystem processes in natural systems is comparatively unknown, but important for the intersection of vertebrate food webs, belowground processes, and ecological heterogeneity. 2. We assessed in situ soil and plant responses to wolf‐killed mammal carrion in Yellowstone National Park, USA. 3. Bison and elk carcasses increased soil respiration and vegetation nutrient concentration and altered bacterial and fungal communities on carcass compared to control plots. The “fingerprints” of soil microbial taxa associated with bison compared to elk carcasses differed considerably and taxa found depended upon abiotic gradients and soil properties. 4. We found evidence that soil microbial community changes associated with carcasses may not be as generalizable as previously thought, which is important for a mechanistic understanding of the links between carrion and soil heterogeneity and potentially for applications in forensic science. 5. This work demonstrates the importance of carrion studies in natural systems. Our findings show that carrion creates distinct ecological patterns that contribute to both above‐ and belowground biological heterogeneity, linking carrion distribution dynamics with soil microbial biodiversity and ecosystem functions.
... Most leopard diet studies have been conducted within a particular landscape type such as rainforest (Hart et al., 1996;Henschel et al., 2005Henschel et al., , 2011Sidhu et al., 2015), mountains (Martins et al., 2011;Norton et al., 1986;Rautenbach, 2010;Rodel et al., 2004;Stuart & Stuart, 1993;Taghdisi et al., 2013), anthropogenic landscapes (Athreya et al., 2016), savannah grasslands (Balme et al., 2017;Kissui, 2008) or arid/semi-arid landscapes (Bothma & Le Riche, 1994;Mondal et al., 2011Mondal et al., , 2012Voigt et al., 2018). Leopard diets differ substantially between the studies as these distinct habitats have different arrays of prey species. ...
Article
en Understanding carnivores’ diet is key to understanding their adaptability in a rapidly changing world. However, studying diet of large carnivores is difficult due to their elusive nature. In this study, we performed DNA metabarcoding analyses of 82 putative leopard scats collected from two distinct, but connected, habitat types (rainforest and grassland) in the Udzungwa Mountains, Tanzania. Two mitochondrial markers were used to identify predator and prey. Metabarcoding confirmed that 60 of the collected scats (73%) originated from leopards, and nineteen mammalian prey DNA sequences were identified to species. Using prey size correction factors for leopards, and covariates on habitat type and prey ecology, we investigated whether differences in leopard dietary composition were detectable between habitats. We found that leopards in grassland consumed a larger mean prey size compared with leopards in rainforest. Small prey (<19 kg) constituted >70% of the biomass consumed by leopards in rainforest, while large prey (≥80 kg) were only eaten in grassland. Arboreal species constituted 50% of the biomass consumed by rainforest leopards. Our results highlight the importance of arboreal species in their diet. From a management perspective, we suggest continued protection of all prey species in the protected areas to prevent human–wildlife conflicts. Résumé fr La compréhension du régime alimentaire des carnivores est essentielle pour comprendre leur adaptabilité dans un monde en évolution rapide. Cependant, l'étude du régime alimentaire des grands carnivores est difficile en raison de leur nature insaisissable. Dans cette étude, nous avons effectué des analyses basées sur le métabarcodage de l'ADN de 82 excréments de léopards présumés, collectés dans deux types d'habitats distincts mais liés (forêt tropicale et prairie) au sein des montagnes Udzungwa, en Tanzanie. Deux marqueurs mitochondriaux ont été utilisés pour identifier les prédateurs et les proies. Le métabarcodage a confirmé que 60 des excréments collectés (73%) provenaient de léopards, et dix‐neuf séquences d'ADN de proies de mammifères ont été identifiées à l’espèce. En utilisant des facteurs de correction de la taille des proies pour les léopards et des covariables sur le type d'habitat et l'écologie des proies, nous avons tenté de déterminer si les différences dans la composition du régime alimentaire des léopards étaient détectables entre les différents habitats. Nous avons constaté que les léopards des prairies consommaient des proies dont la taille moyenne était plus importante que celle des proies consommées par les léopards de la forêt tropicale. Les petites proies (<19 kg) constituaient plus de 70% de la biomasse consommée par les léopards de la forêt tropicale, tandis que les grandes proies (≥ 80 kg) étaient uniquement consommées dans les prairies. Les espèces arboricoles constituaient 50% de la biomasse consommée par les léopards de la forêt tropicale. Nos résultats mettent en évidence l'importance des espèces arboricoles dans leur régime alimentaire. En termes de gestion, nous suggérons de continuer à protéger toutes les espèces de proies dans les zones protégées afin de prévenir les conflits entre les hommes et la faune.
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Kleptoparasitism can be considered as a game theoretical problem and a foraging tactic at the same time, so the aim of this paper is to combine the basic ideas of two research lines: evolutionary game theory and optimal foraging theory. To unify these theories, firstly, we take into account the fact that kleptoparasitism between foragers has two consequences: the interaction takes time and affects the net energy intake of both contestants. This phenomenon is modeled by a matrix game under time constraints. Secondly, we also give freedom to each forager to avoid interactions, since in optimal foraging theory foragers can ignore each food type (we have two prey types: either a prey item in possession of another predator or a free prey individual is discovered). The main question of the present paper is whether the zero-one rule of optimal foraging theory (always or never select a prey type) is valid or not, in the case where foragers interact with each other? In our foraging game we consider predators who engage in contests (contestants) and those who never do (avoiders), and in general those who play a mixture of the two strategies. Here the classical zero-one rule does not hold. Firstly, the pure avoider phenotype is never an ESS. Secondly, the pure contestant can be a strict ESS, but we show this is not necessarily so. Thirdly, we give an example when there is mixed ESS.
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Determining populations of leopards (Panthera pardus) is important for both their conservation and also that of their prey. Camera-trapping has emerged as a powerful and non-invasive tool for studying carnivores in their natural habitats especially for species that are elusive or occur at low densities such as leopards. This thesis presents the Baited-Camera Trapping (BCT) method of censusing leopards, a Zimbabwean conceived design modification of the conventional unbaited setup. This method has been documented to improve capture rates and provide robust and novel data for leopard surveys in savanna environments. This study used single cameras coupled with bait to survey a population of leopards at Malilangwe Wildlife Reserve (MWR), a privately owned medium-sized property in south-eastern Zimbabwe. The objectives of the study were to: (1) conduct a cost-benefit analysis to determine the optimal density and exposure length of baits for censusing leopards at MWR, (2) develop a technique for estimating body dimensions of leopards from camera trap photographs, (3) determine the influence of competing carnivores on the feeding habits of leopards in a savanna ecosystem, and (4) to review the application of the BCT method in comparison with conventional camera trapping. Data were collected from July 2017 to January 2018 and the CAPTURE software was used for population size analysis and the Statistical Package for Social Scientists were used for cost-benefit analyses. Generalized Linear Mixed Effects Modelling, performed using the R statistical software, was used to compare actual and photograph based body measurement data as well as to analyze the influence of competing predators on feeding duration and resting distances of leopards at bait stations. This study estimated the leopard population at MWR at 61 (61-67) individuals and concludes that using BCT stations at a density of 0.24 cameras km-2 km for 9 days is the optimal and cost-effective sampling effort required to provide reliable population statics in semi-arid savannas. The study established that the type of body measurement and the posture of a leopard in a photograph had a significant influence on the accuracy of image-based measurements. Body length measurements taken from the level back-straight forelimb-parallel tail posture were the most accurate [mean error = 2.0 cm (1.5-2.7 cm)] while head-to-tail and tail length measurements and variations from the level back-straight forelimb-parallel tail posture did not provide sufficient accuracy. The findings also showed that the presence of male leopards at feeding locations was associated with shorter feeding durations while lion presence caused feeding leopards to wait longer from bait sites. The thesis provides the first published record of the BCT method outlining a step-by-step procedure for replication by other researchers and a comparative review of the method with traditional survey approaches. The findings in this thesis underscore the ability of BCT method to investigate multiple leopard population ecology questions which enhances its cost-benefit ratio. Furthermore, the method provides new information which can broaden the scope of research and inform management and policy direction. It is recommended that (i) researchers and managers incorporate cost-benefit analysis in their work as this is essential for informing effective application of effort and resources, (ii) researchers take advantage of the BCT method to collect behaviour and morphological data for species that are less understood such as leopards to maximize on the capital investment, (iii) managed wildlife areas that contain leopards consider the uptake of the BCT techniques as a wide encompassing population monitoring option, and (iv) regulatory authorities that supervise hunting operations such as the Zimbabwe Parks and Wildlife Management Authority adopt the BCT technique to enhance their information management portfolios and quota setting for sustainable harvest practises.
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Background & Aim: Carnivores play an important role in maintaining the structural and functional stability of ecosystems. Increasing anthropogenic pressure has resulted in dramatic population decline and habitat degradation of carnivores worldwide. Exploring regional coexistence mechanisms of carnivore populations is important to understand how mammal communities assemble, along with protecting and managing endangered species. This review summarizes the interactions of terrestrial carnivores along three dimensions (i.e., spatial, temporal and dietary), of their ecological niches, analyzing the influence of factors such as body size, prey composition, environmental difference, human disturbance and climate change on the coexistence of carnivores based on more than 100 related studies. Review Results: We put forward the problems to be addressed for current study of carnivore coexistence. This study demonstrates that there is no single theoretical explanation for carnivore coexistence through niche separation. Prey,
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Effective conservation management requires an understanding of the spatiotemporal dynamics driving large carnivore density and resource partitioning. In African ecosystems, reduced prey populations and the loss of competing guild members, most notably lion (Panthera leo), are expected to increase the levels of competition between remaining carnivores. Consequently, intraguild relationships can be altered, potentially increasing the risk of further population decline. Kasungu National Park (KNP), Malawi, is an example of a conservation area that has experienced large-scale reductions in both carnivore and prey populations, leaving a resident large carnivore guild consisting of only leopard (Panthera pardus) and spotted hyena (Crocuta crocuta). Here, we quantify the spatiotemporal dynamics of these two species and their degree of association, using a combination of co-detection modeling, time-to-event analyses, and temporal activity patterns from camera trap data. The detection of leopard and spotted hyena was significantly associated with the detection of preferred prey and competing carnivores, increasing the likelihood of species interaction. Temporal analyses revealed sex-specific differences in temporal activity, with female leopard activity patterns significantly different to those of spotted hyena and male conspecifics. Heightened risk of interaction with interspecific competitors and male conspecifics may have resulted in female leopards adopting temporal avoidance strategies to facilitate coexistence. Female leopard behavioral adaptations increased overall activity levels and diurnal activity rates, with potential consequences for overall fitness and exposure to sources of mortality. As both species are currently found at low densities in KNP, increased risk of competitive interactions, which infer a reduction in fitness, could have significant implications for large carnivore demographics. The protection of remaining prey populations is necessary to mitigate interspecific competition and avoid further alterations to the large carnivore guild.
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Protected areas are the basis for many conservation plans for wildlife. However, they are rarely of an adequate size for the long term survival of populations of large, wide-roaming mammals. In the Maasai Mara, Kenya, communally owned wildlife conservancies have been developed to expand the area available for wildlife. As these continue to develop it is important to ensure that the areas chosen are beneficial to wildlife. Using presence data on cheetah (Acinonyx jubatus), elephant (Loxodonta africana), spotted hyaena (Crocuta crocuta), leopard (Panthera pardus), lion (P. leo) and wild dog (Lycaon pictus), collected through interviews with people living outside the protected areas (n=648), we identify key wildlife areas using false positive site-occupancy modelling. The probabilities of site use were first determined per species based on habitat, protected area distance, river distance and human presence and these were then combined to create a species richness map to highlight key wildlife areas. All six species, except hyaena, showed a preference for a certain habitat type and all avoided human presence. Leopard, elephant, lion and wild dog preferred sites closer to rivers. Hyaena showed minimal influence by distance from protected areas but the other five species all selected for sites closer to the protected areas. The resulting species richness map highlights important areas for conservation efforts, including the expansion of wildlife areas and areas where human development, such as a newly tarmacked road, could have an impact on wildlife.
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1.Although interspecific competition plays a principle role in shaping species behaviour and demography, little is known about the population‐level outcomes of competition between large carnivores, and the mechanisms that facilitate coexistence. 2.We conducted a multi‐landscape analysis of two widely distributed, threatened large carnivore competitors to offer insight into coexistence strategies and assist with species‐level conservation. 3.We evaluated how interference competition affects occupancy, temporal activity and population density of a dominant competitor, the lion (Panthera leo), and its subordinate competitor, the leopard (Panthera pardus). We collected camera‐trap data over three years in ten study sites covering 5,070 km2. We used multispecies occupancy modelling to assess spatial responses in varying environmental and prey conditions and competitor presence, and examined temporal overlap and the relationship between lion and leopard densities across sites and years. 4.Results showed that both lion and leopard occupancy was independent of – rather than conditional on – their competitor's presence across all environmental covariates. Marginal occupancy probability for leopard was higher in areas with more bushy, ‘hideable’ habitat, human (tourist) activity and topographic ruggedness, whereas lion occupancy decreased with increasing hideable habitat and increased with higher abundance of very large prey. Temporal overlap was high between carnivores and there was no detectable relationship between species densities. 5.Lions pose a threat to the survival of individual leopards, but they exerted no tractable influence on leopard spatial or temporal dynamics. Furthermore, lions did not appear to suppress leopard populations, suggesting that intraguild competitors can coexist in the same areas without population decline. Aligned conservation strategies that promote functioning ecosystems, rather than target individual species, are therefore advised to achieve cost‐ and space‐effective conservation. This article is protected by copyright. All rights reserved.
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Carnivore kill frequency is a fundamental part of predator–prey interactions, which are important shapers of ecosystems. Current field kill frequency data are rare and existing models are insufficiently adapted to carnivore functional groups. We developed a kill frequency model accounting for carnivore mass, prey mass, pack size, partial consumption of prey and carnivore gut capacity. Two main carnivore functional groups, small prey‐feeders versus large prey‐feeders, were established based on the relationship between stomach capacity (C) and pack corrected prey mass (iMprey). Although the majority of small prey‐feeders is below, and of large prey‐feeders above a body mass of 10–20 kg, both occur across the whole body size spectrum, indicating that the dichotomy is rather linked to body size‐related ecology than physiology. The model predicts a negative relationship between predator size and kill frequency for large prey‐feeders. However, for small prey‐feeders, this negative relationship was absent. When comparing carnivore prey requirements to estimated stomach capacity, small carnivores may have to eat to their full capacity repeatedly per day, requiring fast digestion and gut clearance. Large carnivores do not necessarily have to eat to full gastric capacity per day, or do not need to eat every day, which in turn reduces kill frequencies or drives other ecological processes such as scavenging, kleptoparasitism, and partial carcass consumption. Where ecological conditions allow, large prey‐feeding appears attractive for carnivores, which can thus reduce activities related to hunting. This is particularly so for large carnivores, who can achieve distinct reductions in hunting activity due to their relatively large gut capacity. This article is protected by copyright. All rights reserved.
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Decline in global carnivore populations has led to increased demand for assessment of carnivore densities in understudied habitats. Spatial capture-recapture (SCR) is used increasingly to estimate species densities, where individuals are often identified from their unique pelage patterns. However, uncertainty in bilateral individual identification can lead to the omission of capture data and reduce the precision of results. The recent development of the two-flank spatial partial identity model (SPIM) offers a cost-effective approach, which can reduce uncertainty in individual identity assignment and provide robust density estimates. We conducted camera trap surveys annually between 2016 and 2018 in Kasungu National Park, Malawi, a primary miombo woodland and a habitat lacking baseline data on carnivore densities. We used SPIM to estimate density for leopard (Panthera pardus) and spotted hyaena (Crocuta crocuta) and compared estimates with conventional SCR methods. Density estimates were low across survey years, when compared to estimates from sub-Saharan Africa, for both leopard (1.9 AE 0.19 SD adults/100 km 2) and spotted hyaena (1.15 AE 0.42 SD adults/100 km 2). Estimates from SPIM improved precision compared with analytical alternatives. Lion (Panthera leo) and wild dog (Lycaon pictus) were absent from the 2016 survey, but lone dispersers were recorded in 2017 and 2018, and both species appear limited to transient individuals from within the wider transfrontier conservation area. Low densities may reflect low carrying capacity in miombo woodlands or be a result of reduced prey availability from intensive poaching. We provide the first leopard density estimates from Malawi and a miombo woodland habitat, whilst demonstrating that SPIM is beneficial for density estimation in surveys where only one camera trap per location is deployed. The low density of large carnivores requires urgent management to reduce the loss of the carnivore guild in Kasungu National Park and across the wider transfrontier landscape.
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Individual specialisation, when individuals exploit only a subset of resources utilised by the population, is a widespread phenomenon. It provides the basis for evolutionary diversification and can impact population and community dynamics. Both phenotypic traits and environmental conditions are predicted to influence individual specialisation; however, its adaptive consequences are poorly understood, particularly among large mammalian carnivores that play an important role in shaping ecosystems. We used observations of 2960 kills made by 49 leopards Panthera pardus in the Sabi Sand Game Reserve, South Africa, to quantify the magnitude of individual dietary specialisation in a solitary large carnivore, and to examine the proximate and ultimate drivers of this behaviour. We found evidence of individual specialisation in leopard diet, with respect to both the species and size of prey killed. Males tended to be more specialised than females, likely because they could access a wider range of prey due to larger body size. Similarly, individuals that encountered a greater diversity of prey tended to be more specialised. Our results confirmed that ecological opportunity was a key determinant of individual specialisation; however, contrary to predictions, per capita resource availability (and by extension, intraspecific competition) did not affect the degree of specialisation exhibited by individuals. Surprisingly, dietary specialisation appeared to disadvantage male leopards. Specialist males overlapped with fewer resident females, had fewer cubs born on their home ranges, and had fewer cubs survive to independence on their home ranges than generalist males. This may have resulted from the high degree of environmental stochasticity experienced during our study, as dietary specialisation is expected to advantage individuals more during periods of resource predictability. In summary, we showed that a species usually considered to be a dietary generalist was in fact a heterogenous collection of specialist and generalist individuals. Individual specialisation is typically assumed to be maintained by disruptive and/or fluctuating selection; hence, the somewhat paradoxical coexistence of both in the same population might be explained by a dynamic evolutionary equilibrium that exists between specialists and generalists, in which each benefit under different conditions.
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In Namibia, leopards (Panthera pardus) are widely distributed, used commercially as trophy animals and are often persecuted for perceived or real predation on livestock and valuable game species outside protected areas. Therefore, leopard populations living in protected areas might be important source populations and for maintaining connectivity. Little data on their population sizes and densities are available from the northern part of the country, particularly from protected areas. Here, we estimated leopard densities using a spatial capture-recapture approach in northern Namibia: (i) the Khaudum National Park (KNP) in northeast Namibia with an annual average rainfall of 450 mm and (ii) the Lower Hoanib River (LHR) in northwest Namibia with an annual average rainfall of 25 mm. With an effort of 2430 and 2074 camera trap nights in the KNP and LHR, respectively, 11 adult female and six adult male leopards were identified in the KNP, whilst only one adult female leopard was detected once in the LHR. For the KNP, a maximum likelihood approach (using the package SECR) revealed a density estimate of 2.74 leopards/100 km 2 , whereas a Bayesian approach (using the package SPACECAP) revealed a density estimate of 1.83 leopards/100 km 2. For the LHR, no density estimate could be determined and it is suggested that the leopard density in such an arid environment is low. These are the first leopard density estimates based on camera trap surveys provided for these protected areas and thus of importance for further monitoring programs to understand leopard population dynamics. We discuss our findings with current habitat changes and conservation measures in both study areas.
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Spatial partitioning in ecological communities has predominantly been described in two dimensions, yet habitat is complex and 3D. Complex space use mediates community structure and interaction strength by expanding spatial, temporal, and dietary dimensions. Vertical stratification of resources provides opportunities for novel specializations, creating a 3D niche. Competition and predation are mediated by 3D space use, as individuals use the vertical axis to access prey, flee predators, or avoid competitors. The 3D niche is important for long-term conservation strategies as species must navigate tradeoffs in habitat use between strata-specific threats and suboptimal habitat patches. Ultimately, elucidating the 3D niche has implications for protected area management and corridor design that directly influence species persistence and ecosystem function in a rapidly changing world.
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Mammalian carnivores may be important agents of prehistoric bone accumulations. Taphonomic analyses of bone assemblages used for specific assignment usually include information on feeding, breeding, denning and even defecating ecology of extant species. Here, we review literature for the Hyaenidae, Felidae and Canidae families of carnivores, focusing on the ecological and behavioural traits that are commonly used as criteria to assign bone accumulations to specific carnivores, and whether these correspond to the present behaviour and ecology of extant species. We found a total of 93 records where 12 species (9 extant species) of these families were considered as bone accumulators in archae-ozoological sites. Hyaenidae was the group most often cited, followed by Felidae and Canidae. Crocuta crocuta was by far the species most often cited as a bone accumulator. Most bone deposits assigned to carnivores (84.9%) were found in underground cavities, and to a lesser extent in non-cave deposits (15.1%). The use assigned to the sites was mainly as a den (29.5%) or breeding den (29.5%), followed by prey depot (16.2%), feeding shelter (12.4%), and to a lesser extent a hunting place (7.6%), with some remarkable differences among families. Coprolites were also found in 53.8% of cases. The behaviour of present hyenas may be similar to that of prehistoric ones as they commonly use underground dens, defecate inside of them and frequently accumulate prey remains. On the other hand, even though present canids are more often recorded than felids using underground dens and accumulating prey, the latter are more often recorded as prehistoric bone accumulators than the former. The behaviour of only one present species of canid (V. vulpes) and other a felid (P. pardus) matches the one presumed for prehistoric individuals of such species in relation to bone and scat accumulation. The role of the remaining species as bone and scat accumulator agents in prehistoric sites remains questionable due to differences in their present behaviour. Therefore, many assignments of bone accumulation to specific carnivores are based on assumptions, which did not coincide with the present natural history of the species. Our review also highlights the absence of records of small species as prehistoric bone accumulators.
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Quantification of fine-scale movement, performance, and energetics of hunting by large carnivores is critical for understanding the physiological underpinnings of trophic interactions. This is particularly challenging for wide-ranging terrestrial canid and felid predators, which can each affect ecosystem structure through distinct hunting modes. To compare free-ranging pursuit and escape performance from group-hunting and solitary predators in unprecedented detail, we calibrated and deployed accelerometer-GPS collars during predator-prey chase sequences using packs of hound dogs ( Canis lupus familiaris , 26 kg, n = 4–5 per chase) pursuing simultaneously instrumented solitary pumas ( Puma concolor , 60 kg, n = 2). We then reconstructed chase paths, speed and turning angle profiles, and energy demands for hounds and pumas to examine performance and physiological constraints associated with cursorial and cryptic hunting modes, respectively. Interaction dynamics revealed how pumas successfully utilized terrain (e.g., fleeing up steep, wooded hillsides) as well as evasive maneuvers (e.g., jumping into trees, running in figure-8 patterns) to increase their escape distance from the overall faster hounds (avg. 2.3× faster). These adaptive strategies were essential to evasion in light of the mean 1.6× higher mass-specific energetic costs of the chase for pumas compared to hounds (mean: 0.76 vs. 1.29 kJ kg ⁻¹ min ⁻¹ , respectively). On an instantaneous basis, escapes were more costly for pumas, requiring exercise at ≥90% of predicted $\dot {\mathrm{V }}{\mathrm{O}}_{2\mathrm{MAX}}$ and consuming as much energy per minute as approximately 5 min of active hunting. Our results demonstrate the marked investment of energy for evasion by a large, solitary carnivore and the advantage of dynamic maneuvers to postpone being overtaken by group-hunting canids.
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Caching of animal remains is common among carnivorous species of all sizes, yet the effects of caching on larger prey are unstudied. We conducted a summer field experiment designed to test the effects of simulated mountain lion (Puma concolor) caching on mass loss, relative temperature, and odor dissemination of 9 prey-like carcasses. We deployed all but one of the carcasses in pairs, with one of each pair exposed and the other shaded and shallowly buried (cached). Caching substantially reduced wastage during dry and hot (drought) but not wet and cool (monsoon) periods, and it also reduced temperature and discernable odor to some degree during both seasons. These results are consistent with the hypotheses that caching serves to both reduce competition from arthropods and microbes and reduce odds of detection by larger vertebrates such as bears (Ursus spp.), wolves (Canis lupus), or other lions.
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Aggression by top predators can create a “landscape of fear” in which subordinate predators restrict their activity to low-risk areas or times of day. At large spatial or temporal scales, this can result in the costly loss of access to resources. However, fine-scale reactive avoidance may minimize the risk of aggressive encounters for subordinate predators while maintaining access to resources, thereby providing a mechanism for coexistence. We investigated fine-scale spatiotemporal avoidance in a guild of African predators characterized by intense interference competition. Vulnerable to food stealing and direct killing, cheetahs are expected to avoid both larger predators; hyenas are expected to avoid lions. We deployed a grid of 225 camera traps across 1,125 km2 in Serengeti National Park, Tanzania, to evaluate concurrent patterns of habitat use by lions, hyenas, cheetahs, and their primary prey. We used hurdle models to evaluate whether smaller species avoided areas preferred by larger species, and we used time-to-event models to evaluate fine-scale temporal avoidance in the hours immediately surrounding top predator activity. We found no evidence of long-term displacement of subordinate species, even at fine spatial scales. Instead, hyenas and cheetahs were positively associated with lions except in areas with exceptionally high lion use. Hyenas and lions appeared to actively track each, while cheetahs appear to maintain long-term access to sites with high lion use by actively avoiding those areas just in the hours immediately following lion activity. Our results suggest that cheetahs are able to use patches of preferred habitat by avoiding lions on a moment-to-moment basis. Such fine-scale temporal avoidance is likely to be less costly than long-term avoidance of preferred areas: This may help explain why cheetahs are able to coexist with lions despite high rates of lion-inflicted mortality, and highlights reactive avoidance as a general mechanism for predator coexistence.
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The study of competition and coexistence among similar interacting species has long been considered a cornerstone in evolutionary and community ecology. However, understanding coexistence remains a challenge. Using two similar and sympatric competing large carnivores, Eurasian lynx and wolverines, we tested the hypotheses that tracking among heterospecifics and reactive responses to potential risk decreases the probability of an agonistic encounter when predators access shared food resources, thus facilitating coexistence. Lynx and wolverines actively avoided each other, with the degree of avoidance being greater for simultaneous than time-delayed predator locations. Wolverines reacted to the presence of lynx at relatively short distances (mean: 383 m). In general, lynx stayed longer, and were more stationary, around reindeer carcasses than wolverines. However, when both predators were present at the same time around a carcass, lynx shortened their visits, while wolverine behavior did not change. Our results support the idea that risk avoidance is a reactive, rather than a predictive, process. Since wolverines have adapted to coexist with lynx, exploiting lynx-killed reindeer carcasses while avoiding potential encounters, the combined presence of both predators may reduce wolverine kill rate and thus the total impact of these predators on semi-domestic reindeer in Scandinavia. Consequently, population management directed at lynx may affect wolverine populations and human-wolverine conflicts.
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Leopards (Panthera pardus) are the most widely distributed wild felid in the world, living sympatrically with numerous competitively dominant species in various large carnivore guilds. Leopards generally feed on small to medium-sized ungulates and risk kleptoparasitism from intraguild competitors. One unique, adaptive response to intraguild competition is arboreal caching ('hoisting') by leopards. Hoisting behaviour is thought to vary in frequency among individuals, populations, and between sexes. In our study, leopards fed primarily (85% of observed kills) on impala (Aepyceros melampus), which is the most locally abundant potential prey species. Although we did not detect statistical differences in hoisting rates among individuals within each sex, our results confirmed moderate hoisting rates of 35% in an area with medium densities of lions (Panthera leo), spotted hyaenas (Crocuta crocuta) and African wild dogs (Lycaon pictus). Sex differences in prey hoisting were related to size, and males as the larger sex, hoisted kills (47.7%, n = 44) more often than overlapping females (27%, n = 62). Female leopards hoisted 25-33% of detected kills irrespective of prey size, season, habitat type, presence of dependent cubs, or location in relation to high competitor density areas ('competitor hotspots'). Consistent with this, males hoisted kills more often where visibility and, therefore, probability of detection was highest. We conclude that hoisting is likely correlated with energetic costs, seasonally increased detection by competitors, and direct competitor interactions rather than the potential risk of conflicts within generalized areas of high competitor use ('hotspots').
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Carnivore attacks on livestock are a primary driver of human–carnivore conflict and carnivore decline globally. Livestock depredation is particularly threatening to carnivore conservation in Central India, a priority landscape and stronghold for the endangered tiger. To strengthen the effectiveness of conflict mitigation strategies, we examined the spatial and temporal patterns and physical characteristics of livestock depredation in Kanha Tiger Reserve. We combined livestock compensation historical records (2001–2009) with ground surveys (2011–2012) and carnivore scat to identify when and where livestock species were most vulnerable. Between 400 and 600 livestock were reported for financial compensation each year, and most (91–95 %) were successfully reimbursed. Tigers and leopards were responsible for nearly all livestock losses and most often killed in the afternoon and early evening. Cattle and buffalo were most at risk in dense forests away from villages and roads, whereas goats were most often killed in open vegetation near villages. A spatial predation risk model for cattle revealed high-risk hotspots around the core zone boundary, confirming the significant risks to livestock grazing illegally in the core. Such ecological insights on carnivore–livestock interactions may help improve species-specific livestock husbandry for minimizing livestock losses and enabling coexistence between people and carnivores.
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Scatter hoarding is a common food-hoarding strategy of many granivores, known to significantly contribute to seed dispersal. Models of scatter hoarding typically hold that the spatial distribution of the scatter hoards made by birds and mammals result from a trade-off between the energetic cost of spacing caches and the increased risk of pilferage as cache densities increase. Here, we present evidence from 3 field experiments that eastern gray squirrels (Sciurus carolinensis) rely on an alternative strategy in which preferred food items are stored in open habitats, beyond tree crowns, where the probability of predation is higher but the risk of cache pilferage is also reduced (hereafter the habitat structure hypothesis). In the first 2 experiments, squirrels cached larger, more profitable acorns at significantly greater distances from canopy edges than smaller acorns, which were more often cached under trees. In the third experiment, in which we tested the effects of both tree canopy cover and cache density on pilferage rates, we found no effect of density on pilferage rates but significantly higher pilferage rates under canopy cover. Our results indicate that habitat heterogeneity, potentially as it relates to predation risks, influences the distribution of scatter-hoarded seeds, with more profitable seeds placed in more open vegetation beyond the shadow of parent plants. We suggest that this behavior may reflect a general strategy of other scatter-hoarding mammals and birds that likely increases the probability of dispersal and establishment of seeds.
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One hypothesis for how carnivores with overlapping ecology coexist in natural systems is through heterogeneous competition land-scapes, in which subordinates utilize "competition refuges" to mitigate risks associated with dominant competitors. We tested for the effects of American black bear (Ursus americanus) kleptoparasitism on puma (Puma concolor) foraging in 2 systems in North America. We also tested whether partial prey consumption exhibited by pumas in the presence of bears was better explained by rules of opti-mal foraging or by kleptoparasitism by black bears, and whether pumas utilized spatial competition refuges to mitigate competition with bears over carcass remains. Puma kill rates in ungulates/wk were equivalent across study systems, but 48% greater in the bear season than the no-bear season. Our analyses of handling time did not support the notion that partial prey consumption exhibited by pumas followed patterns of optimal foraging. Rather, puma handling time and prey consumption were better explained by the pres-ence of bears. Surprisingly, pumas did not utilize spatial competition refuges to mitigate competition with black bears, and instead our results suggested they increase their kill rates to compensate for losses. Our results linking high seasonal kill rates of a top predator with kleptoparasitism by a dominant competitor provide strong evidence that the effects of predation can only be understood within a community framework. In particular, we propose that future predation studies should differentiate between relative contributions of predators and competitors on prey dynamics. Further, our results suggest kleptoparasites may indirectly impact prey populations through their effects on top predators.
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Pumas (Puma concolor) live in diverse, often rugged, complex habitats. The energy they expend for hunting must account for this complexity but is difficult to measure for this and other large, cryptic carnivores. We developed and deployed a physiological SMART (species movement, acceleration, and radio tracking) collar that used accelerometry to continuously monitor energetics, movements, and behavior of free-ranging pumas. This felid species displayed marked individuality in predatory activities, ranging from low-cost sit-and-wait behaviors to constant movements with energetic costs averaging 2.3 times those predicted for running mammals. Pumas reduce these costs by remaining cryptic and precisely matching maximum pouncing force (overall dynamic body acceleration = 5.3 to 16.1g) to prey size. Such instantaneous energetics help to explain why most felids stalk and pounce, and their analysis represents a powerful approach for accurately forecasting resource demands required for survival by large, mobile predators.
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Many animals hoard seeds for later consumption and establish seed caches that are often located at sites with specific environmental characteristics. One explanation for the selection of non-random caching locations is the avoidance of pilferage by other animals. Another possible hypothesis is that animals choose locations that hamper the perishability of stored food, allowing the consumption of unspoiled food items over long time periods.We examined seed perishability and pilferage avoidance as potential drivers for caching behaviour of spotted nutcrackers (Nucifraga caryocatactes) in the Swiss Alps where the birds are specialized on caching seeds of Swiss stone pine (Pinus cembra).We used seedling establishment as an inverse measure of seed perishability, as established seedlings cannot longer be consumed by nutcrackers. We recorded the environmental conditions (i.e. canopy openness and soil moisture) of seed caching, seedling establishment and pilferage sites.Our results show that sites of seed caching and seedling establishment had opposed microenvironmental conditions. Canopy openness and soil moisture were negatively related to seed caching but positively related to seedling establishment, i.e. nutcrackers cached seeds preferentially at sites where seed perishability was low. We found no effects of environmental factors on cache pilferage, i.e. neither canopy openness nor soil moisture had significant effects on pilferage rates. We thus could not relate caching behaviour to pilferage avoidance.Our study highlights the importance of seed perishability as a mechanism for seed-caching behaviour, which should be considered in future studies. Our findings could have important implications for the regeneration of plants whose seeds are dispersed by seed-caching animals, as the potential of seedlings to establish may strongly decrease if animals cache seeds at sites that favour seed perishability rather than seedling establishment.
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1.Long-term studies on large felids are rare and yet they yield data essential to understanding the behaviour of species and the factors that facilitate their conservation. 2.We used the most extensive data set so far compiled on leopards Panthera pardus to establish baseline reproductive parameters for females and to determine the demographic and environmental factors that affect their lifetime reproductive success. 3.We used comprehensive sightings reports and photographs from ecotourism lodges in the Sabi Sand Game Reserve, South Africa, to reconstruct life histories for 44 female leopards that gave birth to 172 litters over a 32-year period. 4.Leopards appeared to exhibit a birth pulse; most litters were born in the wet season, particularly in December. Mean age at first parturition (n = 26, mean ± standard error = 46 ± 2 months, range = 33–62) was older than previously recorded, possibly due to elevated intraspecific competition. Average litter size was 1.9 ± 0.1 (n = 140, range = 1–3) and declined with maternal age. Age of litters at independence (n = 52, 19 ± 1 months, range = 9–31) was inversely related to prey abundance but did not affect the likelihood of recruitment of offspring. Interbirth intervals differed following successful litters (in which at least one cub survived to independence; n = 55, 25 ± 1 months, range = 14–39) and unsuccessful litters (n = 46, 11 ± 1 months, range = 4–36), as did the time taken to replace litters. 5.Variation in lifetime reproductive success was influenced mainly by differences in cub survival, which was related to maternal age and vulnerability to infanticide. Cub survival (37%) declined as females got older, perhaps because mothers relinquished portions of their home ranges to philopatric daughters. Male leopards were responsible for many (40%) cub deaths and females appeared to adopt severalstrategies to counter the risk of infanticide, including paternity confusion and displaying a period of reduced fertility immediately after a resident male was replaced. 6.Our results suggest that the reproductive success of female leopards is regulated primarily by top-down processes. This should be taken into account in management decisions, particularly when managers are considering the implementation of invasive activities such as legal trophy hunting.
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Predation and scavenging have been classically understood as independent processes, with predator–prey interactions and scavenger–carrion relationships occurring separately. However, the mere recognition that most predators also scavenge at variable rates, which has been traditionally ignored in food-web and community ecology, leads to a number of emergent interaction routes linking predation and scavenging. The general goal of this review is to draw attention to the main inter-specific interactions connecting predators (particularly, large mammalian carnivores), their live prey (mainly ungulates), vultures and carrion production in terrestrial assemblages of vertebrates. Overall, we report an intricate network of both direct (competition, facilitation) and indirect (hyperpredation, hypopredation) processes, and provide a conceptual framework for the future development of this promising topic in ecological, evolutionary and biodiversity conservation research. The classic view that scavenging does not affect the population dynamics of consumed organisms is questioned, as multiple indirect top-down effects emerge when considering carrion and its facultative consumption by predators as fundamental and dynamic components of food webs. Stimulating although challenging research opportunities arise from the study of the interactions among living and detrital or non-living resource pools in food webs.
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