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Prey preferences of the spotted hyaena (Crocuta crocuta) and degree of dietary overlap with the lion (Panthera leo)
Journal of Zoology
Abstract
Spotted hyaenas Crocuta crocuta were once considered mere scavengers; however, detailed research revealed that they are very efficient predators. Information on what spotted hyaenas actually prefer to prey on and what they avoid is lacking, as well as the factors that influence prey selection. Data from 14 published and one unpublished study from six countries throughout the distribution of the spotted hyaena were used to determine which prey species were preferred and which were avoided using Jacobs' index. The mean of these values for each species was used as the dependent variable in multiple regression, with prey abundance and prey body mass as predictive variables. In stark contrast to the rest of Africa's large predator guild, spotted hyaenas do not preferentially prey on any species. Also surprisingly, only buffalo, giraffe and plains zebra are significantly avoided. Spotted hyaena most prefer prey within a body mass range of 56–182 kg, with a mode of 102 kg. The dietary niche breadth of the spotted hyaena is similar to that of the lion Panthera leo, and the two species have a 58.6% actual prey species overlap and a 68.8% preferred prey species overlap. These results highlight the flexible and unselective nature of spotted hyaena predation and are probably a reason for the species' success throughout its range, despite a large degree of dietary overlap with lions.
... Thus, sites where large felids played a relevant role, such as the Orce sites, do not show a high abundance of small prey because they were not frequently preyed upon by these predators. In present-day African ecosystems, lions (Panthera leo) and spotted hyenas (Crocuta crocuta) prey on different species, most of which weigh less than 600 kg [102,103] and their prey selection overlaps significantly [104]. In Kruger National Park, for example, spotted hyenas prey primarily on wildebeest (Connochaetes taurinus, 163 kg), buffalo (Syncerus caffer, 520 kg), Burchell's zebra (Equus quagga burchellii, 220 kg), greater kudu (Tragelaphus strepsiceros, 170 kg) and impala (Aepyceros melampus, 44 kg) [105] and for lions, buffalo, wildebeest and zebra, with a distinction between males and females, with males preferring buffalo and females wildebeest and zebra [106]. ...
... In Serengeti National Park, total CC is approximately 2875 kg/km 2 *year and 2158 kg/km 2 *year excluding megafauna, with these species accounting for 82% of the CC [31]. Spotted hyenas have a preferred prey range of 56 to 182 kg, with the mode at 102 kg (Hayward 2006). This range may be slightly higher for lions, with wildebeest being the preferred prey in parts of South Africa and East Africa [27]. ...
... This range may be slightly higher for lions, with wildebeest being the preferred prey in parts of South Africa and East Africa [27]. Nevertheless, both spotted hyenas and lions may prey on young individuals of megafauna, such as elephants, giraffes, hippopotamuses and rhinoceroses [116,117], but Hayward [104] argued that hyenas tend to avoid buffalo, plains zebras and giraffes. Given the above, we can generally expect hyenas and lions to hunt prey between 50 and 200 kg, which tend to be the most abundant, with megaherbivores being a less exploited resource, as well as species under 50 kg, except in poor resource conditions, as in the Kalahari [31,[107][108][109][110][111]. ...
The Early Pleistocene sites of Orce in southeastern Spain, including Fuente Nueva-3 (FN3), Barranco León (BL) and Venta Micena (VM), provide important insights into the earliest hominin populations and Late Villafranchian large mammal communities. Dated to approximately 1.4 million years ago, FN3 and BL preserve abundant Oldowan tools, cut marks and a human primary tooth, indicating hominin activity. VM, approximately 1.6 million years old, is an outstanding site because it preserves an exceptionally rich assemblage of large mammals and predates the presence of hominins, providing a context for pre-human conditions in the region. Research suggests that both hominins and giant hyenas were essential to the accumulation of skeletal remains at FN3 and BL, with secondary access to meat resources exploited by saber-toothed felids. This aim of this study aims to correlate the relative abundance of large herbivores at these sites with their estimates of Carrying Capacity (CC) and Total Available Biomass (TAB) using the PSEco model, which incorporates survival and mortality profiles to estimate these parameters in paleoecosystems. Our results show: (i) similarities between quarries VM3 and VM4 and (ii) similarities of these quarries with BL-D (level D), suggesting a similar formation process; (iii) that the role of humans would be secondary in BL-D and FN3-LAL (Lower Archaeological Level), although with a greater human influence in FN3-LAL due to the greater presence of horses and small species; and (iv) that FN3-UAL (Upper Archaeological Level) shows similarities with the expected CC values for FN3/BL, consistent with a natural trap of quicksand scenario, where the large mammal species were trapped according to their abundance and body mass, as there is a greater presence of rhinos and mammoths due to the greater weight per unit area exerted by their legs. Given the usefulness of this approach, we propose to apply it first to sites that have been proposed to function as natural traps.
... The diet of hyenas has received considerable attention in the scientific literature (Kruuk, 1972;Bearder, 1977;Mills, 1990;Yirga et al., 2012). Hyenas are opportunistic feeders and do not have distinct prey species preference (Hayward, 2006). They are highly adaptive feeders; they are hunters of large or medium prey and scavengers of human waste, bones, dung and even anthrax-infested carcasses (Yirga et al., 2012). ...
... Even within the two national parks, Awash and Chebera Churchura, prey remains of domestic species were dominant. This presents a new finding on hyena diets in national parks, as previous studies showed a variety of prey, predominantly wild prey (Honer et al., 2002;Hayward, 2006;Trinkel, 2010). This is probably due to very low densities of natural prey in these national parks, and/or due to scavenging around the houses of park rangers and near villages. ...
... In the present study, frequencies of prey remains of cattle, sheep, donkey and goat were highest, in decreasing order. Unlike other large carnivores, hyenas do not exhibit a preference for any species of prey (Hayward, 2006). The lack of prey preferences might be due to the fact that they are able to meet their food requirements through scavenging (Hayward, 2006). ...
Livestock depredation and scavenging of waste by hyena occurs widely
across Ethiopia. Here we report on an extensive survey of depredation in
10 areas across the country. We found that even hyena from national parks
predominantly feed on anthropogenic waste. We quantified the economic
impact of hyena predation on livestock using semi-structured interviews
with 3,080 randomly selected households. They reported losses of 2,230
domestic animals, 3.9% of their stock or an average annual financial loss
of US $10.3 per household, over the past five years. The diet of hyenas was
assessed in 17 areas across the country, including national parks, by scat
analysis and showed only prey items of domestic origin except in Chebera
Churchura national park where a few items of prey of wild species were
found. Frequencies of prey remains of cattle, sheep, donkey, and goat were
highest in decreasing order. Some hairs in scat originated from depredation,
but most food intake was from waste dumps and slaughterhouses. Survival
of hyenas in Ethiopia is thus largely and widely dependent on management
of livestock conflict and waste. Further research should identify the specific
causes of domestic animal loss and options to mitigate depredation.
... For example, a reduction in the density of African lions (Panthera leo; hereafter lions) could provide a more favourable environment for sympatric species, like spotted hyenas and leopards. Lions and spotted hyenas have been shown to exert considerable pressure on each other mainly through interference competition (Kruuk and Turner 1967;Hayward 2006), and kleptoparasitism. Much of this is owed to their strong degree of dietary overlap, and direct competition over the same food resources (Kruuk and Turner 1967;Hayward 2006). ...
... Lions and spotted hyenas have been shown to exert considerable pressure on each other mainly through interference competition (Kruuk and Turner 1967;Hayward 2006), and kleptoparasitism. Much of this is owed to their strong degree of dietary overlap, and direct competition over the same food resources (Kruuk and Turner 1967;Hayward 2006). Similarly, leopards appear to avoid spotted hyenas temporally in some landscapes, avoiding peak spotted hyena activity periods (Havmøller et al. 2020). ...
... Spotted hyenas are the most successful of Africa's large carnivores, in that they are widespread, and highly adaptable (Mills and Harvey 2001;Hayward 2006). This is due to their generalist diet and ability to access a variety of prey sizes, mainly resultant of both group and solitary foraging (optimal prey range of 56-182 kg). ...
African leopards (Panthera pardus pardus) and spotted hyenas (Crocuta crocuta) are data deficient across much of Africa, and there are only a handful of recent population estimates for these species from Uganda. This has conservation ramifications, as both species are important for wildlife tourism, and leopards are hunted for sport in several regions adjacent to national parks as part of a government-led revenue-sharing scheme to foster increased tolerance of wildlife. We ran a single-season camera-trap survey in each of the northern and southern sections of the Queen Elizabeth Conservation Area (2400 km 2), Uganda's second largest national park. We applied spatially explicit capture-recapture (SECR) models to estimate the population density and abundance of leopards and spotted hyenas in the northern Mweya and Kasenyi plains area, and the southern Ishasha sector. Leopard densities were estimated to be 5.03 (95% Highest Posterior Density, HPD = 2.80-7.63) and 4.31 (95% HPD = 1.95-6.88) individuals/100 km 2 for the north and south of the conservation area, respectively, while spotted hyena densities were 13.44 (95% HPD = 9.01-18.81) and 14.07 individuals/100 km 2 (95% HPD = 8.52-18.54) for the north and south, respectively. Leopard densities were in the middle range of those recorded in the literature, while sex ratios were what would be expected for this polygamous felid. Spotted hyena densities were on the higher end of those recorded for the species using spatially explicit capture-recapture (SECR) methods. Our work provides the first robust population estimate of leopards and spotted hyenas in the Queen Elizabeth Conservation Area of western Uganda.
... Because our framework is prey-centric, we require a prediction of the expected predator mass given an herbivore of body size M C . For larger predators and prey (> 10 5 g) [39][40][41][42][43][44], the expected predator mass given a particular herbivore mass follows roughly E{M P } = v 0 M v1 C , where v 0 = 9.76×10 3 g and v 1 = 0.21 ( Fig. 4b; see Supplementary Materials Appendix 3). Accordingly, larger terrestrial herbivores tend to suffer mortality from proportionately smaller predators, an asymmetry that becomes more pronounced with increasing size [cf. ...
... To obtain an herbivore-centric measure of the expected predator mass given a particular herbivore mass E{M P |M C }, we first compiled the known diets of largebodied predators, including tigers, lions, hyenas, leopards, dhole, wild dogs, and cheetahs [39][40][41][42][43][44]. Because smaller mammalian predators and prey have very differ-ent PPMR relationships than larger-bodied mammalian predators and prey, we here focus exclusively on the predators of large-bodied herbivore prey > 10 5 g. ...
... Because smaller mammalian predators and prey have very differ-ent PPMR relationships than larger-bodied mammalian predators and prey, we here focus exclusively on the predators of large-bodied herbivore prey > 10 5 g. From the mean proportional reliance of predators on largebodied prey [39][40][41][42][43][44], we repeatedly sampled predator dietary distributions to reflect each predator's reliance as a function of prey mass. We introduced variability in predator and prey masses by assuming that body sizes were normally distributed about the expected value with a standard deviation of ±25%, allowing us to obtain a distribution of expected predator diets as a function of prey mass. ...
Energy flow through consumer-resource interactions is largely constrained by body size. Allometric relationships govern the dynamics of populations by impacting rates of reproduction, as well as alternative sources of mortality, which have differential impacts on smaller to larger organisms. Here we derive the timescales associated with four alternative sources of mortality for terrestrial mammals: mortality from starvation, mortality associated with aging, mortality from consumption by predators, and mortality introduced by subsidized harvest. The incorporation of these allometric relationships into a minimal consumer-resource model illuminates central constraints that may contribute to the structure of mammalian communities. Our framework reveals that while starvation largely impacts smaller-bodied species, the allometry of senescence is expected to be more difficult to observe. In contrast, external predation and subsidized harvest primarily influence larger-bodied species. The inclusion of predation mortality reveals mass thresholds for mammalian herbivores, where dynamic instabilities may limit the feasibility of megafaunal populations. Moreover, we show how these thresholds vary with alternative predator-prey mass ratios, a relationship that is little understood within terrestrial systems. Finally, we use our framework to predict the harvest pressure required to induce mass-specific extinctions, which closely align with previous estimates of megafaunal exploitation in both paleontological and historical contexts.
... Spotted hyaenas are flexible in their behaviour, habitat tolerance and preferred prey species (Holekamp and Dloniak 2010). The wide dietary breadth of spotted hyaenas can lead to a high degree of prey overlap with other carnivores, most notably lions (~60% overlap ;Hayward 2006;Hayward and Kerley 2008;Périquet et al. 2014;Comley et al. 2020a). Leopards are the most widespread large carnivore found throughout Africa (du Preez et al. 2014;Swanepoel et al. 2015;Jacobson et al. 2016). ...
... Spotted hyaenas and lions appeared to avoid each other spatially, whereas spotted hyaenas and leopards, and lions and leopards utilised similar areas. Hayward (2006) and Périquet et al. (2014) found that the high dietary overlap between lions and spotted hyaenas can lead to spatial partitioning between them to avoid interference competition. On Madikwe, lions and spotted hyaenas have the highest dietary overlap among the three large carnivores (80%), followed by leopards and spotted hyaenas (64%) and (Hayward and Kerley 2008;Balme et al. 2017b;Havmøller et al. 2020;Honiball et al. 2021), and the consequently reduced chance of interference competition, could negate the need for spatial exclusion between leopards and spotted hyaenas, and leopards and lions. ...
... Leopards and lions did not appear to affect each other spatially, similar to other studies (see Maputla et al. 2015;Miller et al. 2018;Balme et al. 2019;Rafiq et al. 2020). The dietary overlap between lions and leopards (ranging from 35.1% to 58.6% ;Hayward 2006;Hayward and Kerley 2008;Balme et al. 2017b) is less than that of lions and spotted hyaenas, potentially reducing the need for these carnivores to avoid each other in space. 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). ...
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.
... Spotted-hyaenas occur across many different biomes and vegetation densities and do not generally hide kills (Bohm and Höner 2015;Kruuk 1972), and thus may be particularly important in providing carcasses suitable for vultures. Spotted-hyaenas have a preferred wildlife prey weight range of 91-139+ kg (Clements et al. 2014;Hayward 2006;Owen-Smith and Mills 2008) and are able to subdue larger wildlife species (Kruuk 1972;Trinkel 2009b) such as wildlife in excess of 300 kg (Cooper, Holekamp, and Smale 1999;Kruuk 1972). Several studies highlight their aptitude for hunting over facultative scavenging (Cooper, Holekamp, and Smale 1999;Gasaway, Mossestad, and Standers 1991;Holekamp et al. 1997). ...
... Once carcasses fitting the selection criteria were located, the species was identified, and the percent of soft tissue already consumed was estimated. The approximate age and sex of the carcass were identified where possible to enable estimates of the live weight prior to death (Table S3; species-specific age and sex weights obtained from Estes (2012) and Hayward (2006)). The distance of the carcass from the transect being driven was also measured, to enable estimation of carcass density for each property accounting for differences in detectability. ...
Changes in predator guild composition can affect obligate scavengers through facilitation and competition dynamics. In Namibia, declines of spotted‐hyaenas ( Crocuta crocuta ) may influence threatened vultures, either positively, via provisioning carcasses, or negatively, as dominant scavengers competing for carcasses. To examine potential mechanisms of influence of spotted‐hyaenas on vultures, we compared carcass densities, carcass size (live weight estimated by species, age class and sex), and potential visibility of carcasses to vultures between sites with and without spotted‐hyaenas across Namibia. We sampled thirteen private protected areas (PPAs), six with spotted‐hyaenas present and seven where they were absent. Carcass densities were estimated for each PPA using line‐transect sampling, recording a total of fifty‐four carcasses. There were no significant differences in carcass density, carcass size or carcass distance to cover, as a proxy for visibility, between areas with and without spotted‐hyaenas. These results do not indicate mechanisms of either strong facilitation or strong competition with vultures, suggesting that spotted‐hyaena activity on these Namibian reserves is not detrimental to vultures, but equally may not result in increased carcass availability.
... Spotted hyenas can hunt individually or cooperatively, or scavenge to meet their dietary needs [68]. In addition, spotted hyenas can prey on livestock exceeding their own body mass, a behavior attributed to their lack of a specifc prey preference [69,70]. ...
... In addition, seasonal migrations of wild prey species may draw predators into communal areas during the dry season, leading to more frequent interactions with livestock and humans. Similar seasonal patterns in livestock depredation by spotted hyenas, leopards, and lions have been documented in other studies [60,69,70,73,75,95,99]. Te lack of seasonal variation in leopard predation suggests that leopards may not adjust their livestock attacks in response to migratory prey. ...
Resolving human–carnivore conflict is crucial for the sustainable coexistence of humans and wildlife. Achieving this, however, requires a comprehensive understanding of the causes and complexities associated with the conflict. This study aimed to assess the nature, underlying causes and costs of human–carnivore conflict, as well as the conservation challenges and potential mitigation measures in and around Kafta Sheraro National Park, Ethiopia. In 2020, we conducted 210 questionnaire interviews with villagers surrounding the park. The surveys gathered data on respondents’ reports of carnivore depredation, the extent and patterns of human–carnivore conflicts, incidents of carnivore killings, the economic impact of livestock depredation, threats to wildlife conservation, potential traditional mitigation measures for human–carnivore conflicts, and their socio-economic characteristics. A logistic regression model was used to identify factors contributing to livestock depredation. Seventy-one percent of respondents reported conflict with carnivore species in the 5 years preceding the study. A total of 1390 heads of stock were reported killed by carnivore species, resulting in an economic loss of US$ 170,741. Spotted hyenas (Crocuta crocuta) were responsible for most livestock depredation (37.8%), followed by leopards (Panthera pardus) (29.5%), caracals (Felis caracal) (8%), and common jackals (Canis aureus) (7.3%). Spotted hyenas killed all types of livestock [cattle (Bos taurus), goat (Capra hircus), sheep (Ovis aries), and donkey (Equus asinus)] across all the studied villages, while leopards, caracals, and jackals primarily targeted goats and sheep. Our findings indicated that most livestock depredations occurred at night and during the dry season. Although villagers employed several mitigation measures to prevent carnivore attacks, vigilance and bush fences were reported as the most effective methods. The study also revealed that agricultural land expansion and human settlements are major threats to wildlife conservation. Approximately 75% of surveyed households admitted to retaliatory killings of carnivore species in direct response to livestock losses. The study suggests that effective conflict mitigation and community conservation education should incorporate strategies to promote sustainable wildlife conservation and rural development.
... While there is a rich body of research investigating the prey preference and selection in South African carnivores (e.g. Hayward & Kerley, 2005;Hayward, 2006;Hayward et al., 2006a;Hayward, O'Brian, Hofmeyr & Kerley, 2006b, Clements et al., 2014, little is known about carnivore diets in non-protected areas where predation of livestock would most likely occur (e.g. Forbes, 2011;Humphries, Tharmalingam & Downs, 2016). ...
... In ecosystems with high prey densities, such as the Maasai Mara in Kenya, hyenas kill as much as 95% of the food they eat (Cooper, Holekamp & Smale, 1999). They mostly consume medium to large ungulates weighing up to 350 kg (Hayward, 2006). However, they are also capable of effectively hunting sizeable animals such as giraffe Giraffa camelopardalis giraffa and buffalo (Kruuk, 1972a;Cooper, 1990;East & Hofer, 1993;Holekamp, Smale, Berg & Cooper, 1997). ...
This assessment provides a policy relevant synthesis on the topic of livestock predation and its management in South Africa, as well as recommendations for future research.
See also https://predsa.mandela.ac.za/
... Spotted hyaenas (Crocuta crocuta) (hereinafter referred to as hyaena) are successful, efficient hunters who prey on a wide variety of animal species (Holekamp et al., 1997;Hayward, 2006). Across their range, hyaena diets vary, with individuals selecting prey in accordance with their abundance, both seasonally and geographically (Mills and Hofer, 1998). ...
... Across their range, hyaena diets vary, with individuals selecting prey in accordance with their abundance, both seasonally and geographically (Mills and Hofer, 1998). However, in comparison with other large African carnivores, hyaenas were always regarded as the species with the fewest published dietary studies (Hayward, 2006), especially within Namibia. Fortunately, in the past five years more studies focussing on the diets of these species in Namibia have been conducted, with the most recent published results being that of Fester, Hockings, Van Vuuren and Van Vuuren (Fester et al., 2021), who focusses on the feeding ecology of these hyaenas in the Namib Desert, Namibia and throughout their range, medium sized ungulates predominate in their diets (Périquet et al., 2014), such as impala (Aepyceros melampus) and kudu (Tragelaphus strepsiceros) (Henschel and Skinner, 1990), waterbuck (Kobus ellipsiprymnus) (Holekamp et al., 1997), springbok (Antidorcas marsupialis) (Gasaway et al., 1991) and gemsbok (Oryx gazelle) (Skinner et al., 1992). ...
Although their ecology has been well studied across their range, spotted hyaena (Crocuta crocuta) diets, especially within Namibia, are understudied when compared to other large African carnivores in southern Africa. Large carnivores outside of protected areas often faces persecution by farmers as they are often perceived as being livestock killers and spotted hyaenas on arid farmlands in Namibia does not escape this prejudice. Here we investigated the scat composition of spotted hyaenas outside of protected areas on farmlands in south-western parts of Namibia. A total of 53 scat samples were collected from which 203 hair samples were used for identification purposes. Our results indicate that 69.46% of hairs in scat were of wildlife species whilst 30.54% of hairs were from domesticated species. The asymptote reached also suggested that 53 scat samples were sufficient to describe the spotted hyaeans's diet in this particular area. We suspect that an outbreak of rabies in the study area as well as anthropogenic food sources could have influenced prey consumption. As the majority of prey consumed were of wildlife species (such as kudu Tragelaphus strepsiceros and Hartmann's Mountain zebra Equus zebra hartmannae) it is suggested that should the natural prey base be maintained livestock predation could be mitigated. We recommend that future researchers investigate the diet composition of the only other apex predator in this study area, leopards (Panther pardus) which could describe the prey selection and dietary overlap between these two large carnivores.
... They directly kill 60-95% of the food they eat (Smith and Holekamp, 2019). The dietary niche of spotted hyenas resembles that of lions and the two species have a 58.6% overlap of actual prey species (Hayward, 2006). However, unlike the rest of Africa's large predators, spotted hyenas do not preferentially prey on any species. ...
... However, unlike the rest of Africa's large predators, spotted hyenas do not preferentially prey on any species. Depending on the parts of its range and season, C. crocuta prefers to hunt a variety of small, medium, and large-sized ungulates (Holekamp et al., 1997;Hayward, 2006). The diet of spotted hyenas is very flexible; in some parts of its range hyenas primarily prey on birds and reptiles while in other parts carrion constitutes a substantial part of their food (Mills and Hofer, 1998). ...
Extinct hyenas of the genus Crocuta were typical representatives of the Late Pleistocene mammoth fauna of Eurasia. The geographical distribution of these animals was vast, stretching from the British Isles to the Pacific coast. In this paper, we studied the tooth microwear of Crocuta spelaea from the Prolom 2 grotto in Crimea and the Bukhtarminskaya Cave in eastern Kazakhstan, and C. ultima ussurica from the Geographical Society Cave in the Russian Far East, to provide a comparative analysis of the feeding habits of these two species and to test for ecological differences. We also tested for ontogenic differences in diet through a comparison of juvenile and adult individuals. Our findings show that the microwear patterns of these Late Pleistocene hyenas resemble those of extant spotted hyenas (C. crocuta) but differ from those of extant stripped (Hyaena hyaena) and brown hyenas (Parahyaena brunnea). However, we noted that the number of scratches in the C. ultima ussurica specimens exceeded the variability observed in extant spotted hyenas and instead more closely resembled the microwear signal of typical predators (e.g., lions). Adults and juveniles exhibited similar microwear patterns in terms of the quantity of pits and scratches; however, most juveniles lacked cross scratches, as also observed in most individuals from Prolom 2. This pattern could be related to a higher proportion of meat in the diet of juveniles, without excluding the consumption of bones. Our tooth microwear analysis improves the knowledge of the paleobiology of extinct representatives of the genus Crocuta.
... In addition, when lion was used as dependent variable, plains zebra, African buffalo and livestock were included as independent variables because these are preferred prey species [57][58][59][60][61] . When leopard, and spotted hyena were employed as dependent variables, antelope, zebra, and livestock were included as independent variables, as these are these are some of their preferred prey species [62][63][64] . For all carnivore species (identified to species or not), we used antelopes as an independent variable. ...
... For other species, predator-prey interactions may explain observed less than expected use of the underpasses. Indeed, the use of underpasses by lion, leopard, and hyena, was positively influenced by presence of buffalo, zebra, and antelope, which are their key prey species 59,62,64 . This suggests that predator-prey interactions were important factors in underpass use by the major carnivores in the TCA. ...
Rail and road infrastructure is essential for economic growth and development but can cause a gradual loss in biodiversity and degradation of ecosystem function and services. We assessed the influence of underpass dimensions, fencing, proximity to water and roads, Normalized Difference Vegetation Index (NDVI), presence of other species and livestock on underpass use by large and medium-sized mammals. Results revealed hyenas and leopards used the underpasses more than expected whereas giraffes and antelopes used the underpasses less than expected. Generalized linear mixed-effects models revealed that underpass height influenced use by wildlife, with several species preferring to use taller underpasses. Electric fencing increased underpass use by funneling species towards underpasses, except for elephants and black-backed jackal for which it reduced underpass passage. We also found that the use of underpasses by livestock reduced the probability of use by nearly 50% for wildlife species. Carnivore species were more likely to cross underpasses used by their prey. Buffalo, livestock, and hyenas used underpasses characterized by vegetation with higher NDVI and near water sources while baboons, dik-diks and antelope avoided underpasses with high NDVI. Our findings suggest a need for diverse and comprehensive approaches for mitigating the negative impacts of rail on African wildlife.
... Both species were responsible for 1,102 attacks on cattle, sheep and goats on the edge of Lake Mburo National Park (hereafter LMNP) (spotted hyenas n = 762 or 69%, leopards n = 340 or 31% between January 2009-December 2018, Braczkowski et al., 2020c). Such conflict between these species and pastoralists may have ramifications and at least 19 leopards were killed on the boundary of LMNP in a 4-year period from 2003-2006 Proposal 3), and two hyena clans (each >14 individuals in size) that were regularly viewed by tourists were poisoned in 2007 (R Schenk, 2018, personal communication). However, both species are also important for the wildlife-viewing tourism (Van der Meer, Badza & Ndhlovu, 2016) and in Uganda in 2018 alone, 1,585 people purchased a night game drive permit for leopard viewing in LMNP, equating to US$47,550 in revenue for the Ugandan Wildlife Authority (A Kule, 2018, personal communication). ...
... Unlike leopards, spotted hyenas do not show a negative correlation with lion presence in Africa (Périquet, Fritz & Revilla, 2015) despite intraguild predation and the negative impact that lions can have on hyena reproduction (Watts & Holekamp, 2008). Spotted hyenas may benefit from the presence of lions-and (Hayward, 2006;Davidson et al., 2019). Observed positive correlations in lion and spotted hyena density in many parts of Africa may also be a result of their similar preferred prey base. ...
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.
... In addition, when lion was used as dependent variable, plains zebra, African buffalo and livestock were included as independent variables because these are preferred prey species [44][45][46][47][48] . When leopard, and spotted hyena were employed as dependent variables, antelope, zebra, and livestock were included as independent variables, as these are these are some of their preferred prey species [49][50][51] . For all carnivore species (classi ed and unclassi ed), we used antelopes as an independent variable. ...
... For other species, predator-prey interactions may explain their less than expected use of the underpasses. Indeed, the use of underpasses by lions, leopards, and hyenas, was positively in uenced by presence of buffalo, zebra, and antelope, which are their key prey species 46,49,51 . This suggests that predator-prey interactions were important factors in underpass use by the major carnivores in the TCA. ...
Rail and road infrastructure is essential for economic growth and development but can cause a gradual loss in biodiversity and degradation of ecosystem function and services. We assessed influence of underpass dimensions, fencing, proximity to water and roads, Normalized Difference Vegetation Index (NDVI), presence of other species and livestock on underpass use by large and medium-sized mammals. Results revealed hyenas and leopards used the underpasses more than expected whereas giraffes and antelopes used the underpasses less than expected. Generalized linear mixed effects models (GLMMs) revealed that underpass height influenced their use by wildlife, with several species preferring to use taller underpasses. Electric fencing increased underpass use by funneling species towards underpasses, except for elephants and black-backed jackal for which it reduced underpass passage. GLMMs also revealed that the use of underpasses by livestock reduced the probability of their use by nearly 50% of wildlife species. Carnivore species were more likely to cross underpasses used by their prey. Buffalo, livestock, and hyenas used underpasses with higher NDVI and near water sources while baboons, dik-diks and antelope avoided underpasses with high NDVI. The findings suggest a need for diverse, and comprehensive approach for mitigating the negative impacts of rail on African wildlife.
... They are opportunists, and their prey selection reflects the relative abundance of the available herbivores in the environment (Kruuk, 1972;Bunn, 1983;Höner et al., 2005;Trinkel, 2010). Hyenas effectively prey on ungulates of body mass ranging from 20 to 300 kg (Kruuk, 1972;Cooper et al., 1999;Höner et al., 2005;Hayward, 2006). Larger species are usually available to hyenas as carrion or are occasionally hunted when either young or injured (Cooper et al., 1999;Holekamp and Dloniak, 2010;Bohm and Höner, 2015). ...
The spotted hyena ( Crocuta crocuta ) was an important large carnivore of Pleistocene ecosystems in Africa and Eurasia. Like its modern relatives, this obligate carnivore was adapted to crush and digest bones of its prey and absorb organic matter from bones more efficiently than any other carnivore. This difference in the nutrient resource use between hyenas and most other carnivores led to differences in the isotope flux and variation in the carbon and nitrogen isotopic composition. In our paper, we assess the prey-to-hyena collagen-to-collagen Δ ¹³ C and Δ ¹⁵ N trophic discrimination factor (TDF), a key parameter needed in mixing models used for quantitative reconstruction of diet. We analyzed a Pleistocene hyena den bone accumulation in Perspektywiczna Cave (Poland), with a preserved assemblage of remains containing both hyenas and a wide spectrum of their prey represented by digested bones. With the use of proteomics-based taxonomic identification (ZooMS), we estimated the proportion of prey species in the hyena diet. The modeled collagen-to-collagen TDFs are around +1.6‰ to +1.7‰ for δ ¹³ C and around +3.4‰ to +3.5‰ for δ ¹⁵ N. This study provides new data on the dietary habits of this large carnivore and allows for a more accurate use of isotopic signals in modeling past hyena diets.
... To quantify the relative selection by the wolf towards the wild boar with respect to the other ungulate species, we used the Jacobs selectivity index: (r − p)/(r + p − 2rp), where r is the proportion of each prey in wolf's diet and p is its relative abundance (Bugir et al., 2021;Hayward, 2006;Jacobs, 1974;Upadhyaya et al., 2018). To estimate the relative availability of each ungulate species, we used camera trap data (see paragraph below) to calculate the species-specific, group size-adjusted detection rates (Relative Abundance Index, i.e., RAI individuals , sensu Ferretti et al., 2023). ...
In predator-prey systems, interspecific interactions at the spatial and temporal scale may affect different components of the ecosystem. Despite a growing interest in the topic, little is known about these interactions in many regions, particularly in areas recently recolonized by apex predators, such as the Alps. Using a multidimensional approach, we assessed wolf-wild boar interactions in a recently recolonized protected area of the Western Alps (Gran Paradiso National Park, Italy) by combining camera trapping and wolf scat analyses. First, we quantified the importance of wild boar in the wolf diet by comparing two different periods (October 2018-September 2020 vs. October 2021-September 2022). Second, we evaluated prey selection in 2021-2022, when data on prey availability were collected. Then, we assessed spatiotemporal interactions between the wild boar and the wolf. Finally, we calculated combined spatiotemporal associations between the wolf and the wild boar in comparison to other ungulate prey. We found a remarkable increase in wild boar consumption between the two periods; roe deer and wild boar were consumed in proportion to their availability, while the chamois was positively selected and the red deer and the ibex were less consumed in relation to their availability. Although we found no negative association between wolf and wild boar detection rates, in winter the wild boar limited its temporal overlap with the predator by being less active in sites more frequently used by the wolf, suggesting some spatial modulation of a temporal antipredator response. Among ungulate species, the wild boar had the highest combined spatiotemporal association with the wolf, particularly in autumn-winter. Our results suggest complex interspecific interactions in alpine areas recently recolonized by the wolf.
OPEN ACCESS - https://doi.org/10.1016/j.gecco.2024.e03253
... We examine this transition from the perspective of two predicted behavioural trends: (i) that observations of active hunting should decline at this transitional prey body mass and (ii) that exclusive scavenging should increase around the same transitional prey body mass. We rely on the abundant observations of kill percentages and prey preference among larger-bodied carnivores-including wild dogs, cheetahs, leopards, spotted hyenas, lions and tigers-to assess whether predicted declines in active hunting correlate to observed declines in the field (Hayward, 2006;Hayward et al., 2012;Hayward, Henschel, et al., 2006;Hayward & Kerley, 2005;. In contrast, to compare our predicted transition to observed increases in exclusive scavenging, we must rely on qualitative data documenting increases in scavenging. ...
The costs of foraging can be high while also carrying significant risks, especially for consumers feeding at the top of the food chain.
To mitigate these risks, many predators supplement active hunting with scavenging and kleptoparasitic behaviours, in some cases specializing in these alternative modes of predation.
The factors that drive differential utilization of these tactics from species to species are not well understood.
Here, we use an energetics approach to investigate the survival advantages of hunting, scavenging and kleptoparasitism as a function of predator, prey and potential competitor body sizes for terrestrial mammalian carnivores.
The results of our framework reveal that predator tactics become more diverse closer to starvation, while the deployment of scavenging and kleptoparasitism is strongly constrained by the ratio of predator to prey body size.
Our model accurately predicts a behavioural transition away from hunting towards alternative modes of predation with increasing prey size for predators spanning an order of magnitude in body size, closely matching observational data across a range of species.
We then show that this behavioural boundary follows an allometric power‐law scaling relationship where the predator size scales with an exponent nearing 3/4 with prey size, meaning that this behavioural switch occurs at relatively larger threshold prey body size for larger carnivores.
We suggest that our approach may provide a holistic framework for guiding future observational efforts exploring the diverse array of predator foraging behaviours.
... The spotted hyena (Crocuta crocuta) is a generalist apex predator that preferentially hunts medium-sized wild ungulates or scavenges from carcasses (Kruuk, 1972;Cooper et al., 2001;Hayward, 2006;Trinkel, 2009). Spotted hyenas occasionally predate on livestock animals, consume various anthropogenic food sources, particularly in semi-urban areas, and feed on other carnivores, small mammals, birds, ostrich eggs, and reptiles (Pienaar, 1969, Kruuk, 1972, Mills, 1990, Yirga et al., 2011, Yirga et al., 2012, Hanssen and Cunningham, 2012, Fester et al., 2021. ...
Spotted hyenas (Crocuta crocuta) are known for their flexible foraging behavior and trophic interactions with diverse prey species. However, most studies describing their hunting behavior are focused on predation of large prey. Here we describe the capture rates and behavior of adult and subadult spotted hyenas hunting passerine birds. Hyenas were actively chasing, catching and feeding on red-billed queleas (Quelea quelea), a passerine bird gathering in large flocks at a waterhole in the Etosha National Park, Namibia. In total, we observed 38 successful captures with the average individual capture rate of 21 birds caught per hour. It remains unclear if this is a learned behavior of a single hyena clan or if it could be a widespread interaction across Africa, where the two species co-occur. Although it is unlikely that passerines could make up a substantial part of a spotted hyenas diet regardless of the location, our observations provide a rare documented example of trophic interactions between a large carnivore and small birds. It also represents an addition to a diverse repertoire of hyena foraging behaviors, which confirms their adaptability in obtaining food from non-typical sources.
... According to van Dalum (2013), the type and likelihood of interactions with wildlife differ among communities due to local differences in the abundance and density of the wildlife population. Additional factors include the effectiveness of existing mitigation measures, damage-causing species' habitat use in particular foraging patterns, and predator-livestock size or herbivore crop type preferences (Blackwell et al. 2016;Hayward 2006;Khorozyan & Waltert 2019;Ugarte et al. 2019). Lions were identified by most respondents to contribute to most livestock predation cases in the Wuparo Conservancy. ...
... in both sites demonstrates these species' success in the landscape, owing largely to their adaptability and tolerance of anthropogenic influence (Chinomona et al., 2018;Hayward, 2006). Nonetheless, high elephant poaching in particularly Bico threatens their continued survival (Nieman & Nieman, 2023). ...
Large transboundary protected areas such as the Kavango-Zambezi Transfrontier Conservation Area (KAZA-TFCA), improve mammal conservation by addressing biodiversity threats on regional and international levels, but an adequate understanding of mammal space use and richness within protected areas is critical for effective management planning. This information has however been missing from the Angolan component of the KAZA-TFCA, such as areas of Luengue-Luiana National Park (LLNP), especially compared with neighbouring countries. We used camera trap data from 2021 to 2023 to investigate species richness of medium-and large-sized mammals in two hotspot areas of LLNP, as well as assess the space use patterns of 9 carnivore and 16 herbivore species. A total of 48 species were recorded over 10,781 camera trapping nights, with African elephant (Loxodonta africana) and small antelope (e.g. common duiker Sylvicapra grimmia and steenbok Raphicerus campestris) being the most common. Species composition differed significantly between sites, with 21 species more relatively abundant at Ndunjuru, including cheetah (Acinonyx jubatus) and African wild dog (Lycaon pictus), and eight species more relatively abundant at Bico, including lion (Panthera leo) and hippopotamus (Hippopotamus amphibius). The detection probability of 22 species were affected by one or more ecological (e.g. proximity to rivers) or an-thropogenic (e.g. human settlement) covariate. These results are intended to act as a baseline to monitor conservation effort and aid habitat management in LLNP, as well as similar protected areas within the KAZA-TFCA and elsewhere.
... prey on any species. Depending on the parts of its range and the season, C. crocuta prefers to hunt a variety of small, medium, and largesized ungulates (Hayward, 2006;Holekamp et al., 1997). Evidence of dental microwear suggests that a hypercarnivorous diet was also characteristic of some extinct hyenas (Rivals et al., 2022). ...
Carnivorans are well‐known for their exceptional backbone mobility, which enables them to excel in fast running and long jumping, leading to them being among the most successful predators amongst terrestrial mammals. This study presents the first large‐scale analysis of mobility throughout the presacral region of the vertebral column in carnivorans. The study covers representatives of 6 families, 24 genera and 34 species. We utilized a previously developed osteometry‐based method to calculate available range of motion, quantifying all three directions of intervertebral mobility: sagittal bending (SB), lateral bending (LB), and axial rotation (AR). We observed a strong phylogenetic signal in the structural basis of the vertebral column (vertebral and joint formulae, length proportions of the backbone modules) and an insignificant phylogenetic signal in most characteristics of intervertebral mobility. This indicates that within the existing structure (stabilization of which occurred rather early in different phylogenetic lineages), intervertebral mobility in carnivorans is quite flexible. Our findings reveal that hyenas and canids, which use their jaws to seize prey, are characterized by a noticeably elongated cervical region and significantly higher SB and LB mobility of the cervical joints compared to other carnivorans. In representatives of other carnivoran families, the cervical region is very short, but the flexibility of the neck (both SB and LB) is significantly higher than that of short‐necked odd‐toed and even‐toed ungulates. The lumbar region of the backbone in carnivorans is dorsomobile in the sagittal plane, being on average ~23° more mobile than in artiodactyls and ~38° more mobile than in perissodactyls. However, despite the general dorsomobility, only some representatives of Canidae, Felidae, and Viverridae are superior in lumbar flexibility to the most dorsomobile ungulates. The most dorsomobile artiodactyls are equal or even superior to carnivorans in their ability to engage in dorsal extension during galloping. In contrast, carnivorans are far superior to ungulates in their ability to engage ventral flexion. The cumulative SB in the lumbar region in carnivorans largely depends on the mode of running and hunting. Thus, adaptation to prolonged and enduring pursuit of prey in hyenas is accompanied by markedly reduced SB flexibility in the lumbar region. A more dorsostable run is also a characteristic of the Ursidae, and the peculiar maned wolf. Representatives of Felidae and Canidae have significantly more available SB mobility in the lumbar region. However, they fully engage it only occasionally at key moments of the hunt associated with the direct capture of the prey or when running in a straight line at maximum speed.
... . Behavioral and morphological adaptations can minimize the potential for negative interactions among such competitors (Hunter & Caro, 2008;Sanchez-Barradas & Villalobos, 2020;St-Pierre et al., 2006), by separating their niches along a behavioral dimension (Hutchinson, 1959). For sympatric carnivores, substantial differences in body mass and the subsequent size of prey selected can facilitate coexistence (Hayward, 2006;Karanth & Sunquist, 1995;Kiltie, 1984;Rosenzweig, 1996), whereas subdominant carnivores may occupy an opportunistic niche that maximizes resource use while minimizing competitive encounters (Ramesh et al., 2012). In this study, we investigate whether the coexistence of three large predators in an understudied, but critical, area for conservation could be simply facilitated by differences in activity patterns that also relates to the activity patterns of known key prey species. ...
Predator behaviors influence, and are influenced by, prey and competitor behaviors. Jaguars ( Panthera onca ), pumas ( Puma concolor ), and ocelots ( Leopardus pardalis ) coexist throughout their geographic range as the three largest predators in a multi‐predator community across diverse environments. This study tested for non‐random segregation and overlap in the activity patterns of these felids and their shared prey in the southern buffer zone of the Calakmul Biosphere Reserve, in southern Mexico, using camera traps during February to August 2019. We detected little temporal segregation between the nocturnal activities of jaguars, pumas, and ocelots, although pumas were more active closer to dawn. Jaguars had low activity overlap with species likely to be common prey, whereas ocelots had high overlap with their potential prey. Pumas displayed finer‐scale similarities in activity with species likely to be common prey. In an understudied area of conservation importance, this study shows that temporal segregation is an unlikely mechanism of coexistence. Further research should incorporate spatio‐temporal avoidance and dietary differences to improve our understanding of the mechanisms that drive coexistence between generalist species in a diverse assemblage of threatened felids.
Abstract in Spanish is available with online material.
... Traditionally, diet studies often focused on fundamental ecological topics, such as predator-prey interactions, interspecific competition and facilitation, and resource selection (Carvalho & Gomes, 2004;Mukherjee, Goyal, & Chellam, 1994;Hayward, 2006). More recently, there has been added focus on applying findings to provide management recommendations (Janeiro-Otero et al., 2020;Mengüllüoğlu et al., 2018;Shao et al., 2021). ...
Human-wildlife coexistence requires rigorous, interdisciplinary evidence that promotes effective conservation and management actions. Such evidence-based approaches are conducive to coexistence between local communities and conflict-prone wildlife, such as large carnivores. Yet, little is known about the best path to gaining local community acceptance of large carnivore management strategies, the effects of anthropogenic activity on the persistence of large carnivores, and the way large carnivores interact with threatened wildlife and local communities. This dissertation focuses on the applied ecology and conservation of large carnivores in Ngorongoro Conservation Area (NCA), Tanzania, a multi-use protected area where the local Maasai community lives alongside wildlife. The three objectives of this dissertation are to: (i) identify the best predictors of the acceptance of large carnivore management strategies by local community members, (ii) assess the effects of anthropogenic activity on large carnivore fitness and physiology, and (iii) understand how large carnivores interact with threatened wildlife and local Communities in multi-use protected areas. I use an interdisciplinary approach by combining socio-psychological data from the Maasai community with long-term data on the diet, fitness, and physiology of free-ranging spotted hyenas in the NCA. In Chapter 2, I show that emotions towards and the cultural importance of large carnivores (spotted hyenas (Crocuta crocuta), lions (Panthera leo), and leopards (Panthera pardus)) are much stronger predictors of the acceptance of large carnivore management strategies than livestock depredation is. I also show that depredation by large carnivores is a much smaller source of livestock death than disease and are drought. I demonstrate that spotted hyenas are viewed less positively than both lions and leopards are, though invasive strategies are not accepted for all three species. The results demonstrate that conservation practitioners may focus too much on livestock depredation as the main impediment to coexistence; Rather, they may need to target the respectful fostering of positive emotions through community engagement, while accounting for how different species are viewed. In Chapter 3, I investigate the effect of diurnal pastoralism on fitness and physiology in the Ngorongoro Crater spotted hyena population over a 24-year period by exploiting a natural experiment: two of the Crater's eight spotted hyena clans were exposed to the pastoralism, while the other six were not. By directly measuring the effects of pastoralism on fitness and physiology, I quantify how an anthropogenic activity affects highly conservation-relevant traits. I found no detectable difference in juvenile recruitment (fitness) nor allostatic load (physiology) between the exposed and unexposed clans, indicating that the pastoralism had no major deleterious effect on the spotted hyenas. These results suggest that exposure to anthropogenic activity may be compatible with the persistence of group-living large carnivores, if spatiotemporal overlap between the species' key behaviors and the activity is limited. Finally, in Chapter 4, I use fecal DNA metabarcoding to show that the Ngorongoro Crater spotted hyena population does not regularly consume the black rhinoceros (Diceros bicornis), a Critically Endangered species. I also show that spotted hyenas at least occasionally leave the Crater to forage, based on detections of both Maasai giraffe (Giraffa camelopardalis tippelskirchi) and domestic animals. Furthermore, I found a positive association between spotted hyena age and the propensity to consume domestic animals, which has implications for conflict mitigation. This dissertation sheds light on (i) the most important variables to target when seeking to gain local acceptance for large carnivore management strategies, (ii) the effects of anthropogenic activity on the fitness and physiology of large carnivores, and (iii) how large carnivores interact with threatened wildlife and local communities in multi-use protected areas. Altogether, this dissertation is expected to provide valuable knowledge for the optimization of evidence-based large carnivore conservation and human-carnivore coexistence.
... Of the three large carnivore species captured in this survey, leopard probably display the highest level of habitat plasticity and adaptability to prey items, where prey as large as greater kudu and giraffe calves have been recorded (Hunter 2015), as well as small prey including insects, reptiles, and birds (Hayward et al. 2006). This high level of adaptability has contributed to the maintenance of healthy leopard populations across Africa. ...
The survey was conducted in Angola's Luengue-Luiana National Park to assess large carnivores and their prey. It aimed to evaluate conservation efforts, population trends, and the status of medium- to large-sized mammals. Camera traps were set up in the Panthera Intensive Protection Zone (PIPZ) and the Kwando Intensive Protection Zone (KIPZ), capturing images of 43 mammal species. Leopard, African wild dog, and spotted hyena were identified to individual-level. The survey results offer valuable insights into wildlife distribution and the effectiveness of post-civil war conservation efforts. They will inform future strategies to mitigate human-wildlife conflict and preserve the region's biodiversity.
... According to van Dalum (2013), the type and likelihood of interactions with wildlife differ among communities due to local differences in the abundance and density of the wildlife population. Additional factors include the effectiveness of existing mitigation measures, damage-causing species' habitat use in particular foraging patterns, and predator-livestock size or herbivore crop type preferences (Blackwell et al. 2016;Hayward 2006;Khorozyan & Waltert 2019;Ugarte et al. 2019). Lions were identified by most respondents to contribute to most livestock predation cases in the Wuparo Conservancy. ...
... According to van Dalum (2013), the type and likelihood of interactions with wildlife differ among communities due to local differences in the abundance and density of the wildlife population. Additional factors include the effectiveness of existing mitigation measures, damage-causing species' habitat use in particular foraging patterns, and predator-livestock size or herbivore crop type preferences (Blackwell et al. 2016;Hayward 2006;Khorozyan & Waltert 2019;Ugarte et al. 2019). Lions were identified by most respondents to contribute to most livestock predation cases in the Wuparo Conservancy. ...
... Hyaenas are apex predators capable of hunting prey many times their own size (Holekamp et al., 1997). Furthermore, at our study site and in other areas across their range, hyaenas are known to prey both on warthogs and porcupines (Davidson et al., 2019;Hayward, 2006). Thus, warthogs and porcupines place themselves at risk if they share dens with hyaenas. ...
We have been monitoring spotted hyaena dens with camera traps at our site since 2016. We describe a novel phenomenon: concurrent subterranean den sharing between spotted hyaenas, warthogs, and crested porcupines at a wildlife conservancy in Kenya. We discovered two different hyaena clans that occasionally shared active dens with warthogs and porcupines in 2016, 2017, 2018, and 2019 but not since. We speculate on the reasons why and how this cohabitation arrangement arose and suggest that it might be based on a ‘healthy respect’ for the threats presented by their mutually formidable weaponry. We hope this note will encourage others to describe similar behaviour.
... Conflict with communities are additionally believed to be the primary threat to lions in pastoralist areas, especially because their scavenging behavior make them vulnerable to poisoning (Bauer et al. 2020;Dickman et al. 2014 Leopard, cheetah and African wild dog largely overlap in habitat and prey preference, and their diets usually depend on the availability of locally abundant prey. Of these three species, leopard probably display the highest level of habitat plasticity and adaptability to prey items, where prey as large as greater kudu and giraffe calves have been recorded (Hunter 2015), as well as small prey including insects, reptiles, and birds (Hayward et al. 2006). Similarly, male cheetah coalitions have been documented to take larger prey such as adult blue wildebeest (Caro 1994) as well as smaller prey such as ground-dwelling birds, and African wild dog have been known to prey on anything from hare to zebra and sable antelope (Estes 1991). ...
The results of the 2022 camera trap surveys conducted in the Intensive Protection Zones of Luengue-Luiana National Park in Angola
... As species with small niche breadths are relatively specialised and species with broad niches are considered more generalist species (Carvalho & Cardoso, 2020;Segurado et al., 2011) (Hayward, 2006;Pienaar, 1969). ...
Large carnivore community structure is affected by direct and indirect interactions between intra-guild members. Co-existence between different species within a carnivore guild may occur through diet, habitat or temporal partitioning. Since carnivore species are highly dependent on availability and accessibility of prey, diet partitioning is potentially one of the most important mechanisms in allowing carnivores to co-exist. Intra-guild interactions may vary over time as carnivore prey preference and diet overlap can change due to seasonal changes in resource availability. We conducted scat analysis to compare the seasonal changes in prey preference, diet partitioning and niche breadth of four large carnivore species, namely leopard Panthera pardus, spotted hyena Crocuta crocuta, brown hyena Parahyaena brunnea and wild dog Lycaon pictus in central Tuli, Botswana. Large carnivores in central Tuli display a high dietary overlap, with spotted hyena and brown hyena displaying almost complete dietary overlap and the other carnivore species displaying slightly lower but still significant dietary overlap. Dietary niche breadth for both hyena species was high possibly due to their flexible foraging strategies, including scavenging, while leopard and wild dog showed a relatively low niche breadth, suggesting a more specialised diet. High dietary overlap in central Tuli is possibly explained by the high abundance of prey species in the area thereby reducing competition pressure between carnivore species. Our research highlights the need to assess the influence of diet partitioning in structuring large carnivore communities across multiple study sites, by demonstrating that in prey rich environments, the need for diet partitioning by carnivores to avoid competition may be limited.
... In general, spotted hyaenas are considered to be generalist foragers, feeding on a wide range of available prey (Hayward 2006), and concentrating on the most abundant species of medium to large size. However, they do show some prey selectivity (Cooper 1990, Höner et al. 2002, Wentworth et al. 2011, Périquet et al. 2015b. ...
... Given the intense protection and limited evidence of human activity in the surveyed area, the possibility of populationdegrading activities impacting such an adaptable predator is extremely unlikely. Secondly, spotted hyaenas are flexible in their feeding patterns (Hayward 2006) and are known to feed on ungulates that are available within their accessible environment. Although previous studies have shown that spotted hyaenas prefer prey with a body mass range of 56-182kg, many of their preferred species (e.g. ...
The distribution and status of large carnivores within the Panthera Intensive Protection Zone, Luengue-Luiana National Park, Angola
... Previous studies have calculated prey preference values for each prey type within each study site, then averaged these individual preference values over all study sites for each prey type [26,27]. The problem with this is that prey preference estimates within one study site are relative to the prey types present and thus cannot be directly combined with other study sites. ...
Most natural ecosystems contain animals feeding on many different types of food, but it is difficult to predict what will be eaten when food availabilities change. We present a method that estimates food preference over many study sites, even when number of food types vary widely from site to site. Sampling variation is estimated using bootstrapping. We test the precision and accuracy of this method using computer simulations that show the effects of overall number of food types, number of sites, and proportion of missing prey items per site. Accuracy is greater with fewer missing prey types, more prey types and more sites, and is affected by the number of sites more than the number of prey types. We present a case study using lion (Panthera leo) feeding data and show that preference vs prey size follows a bell-curve. Using just two estimated parameters, this curve can be used as a general way to describe predator feeding patterns. Our method can be used to: test hypotheses about what factors affect prey selection, predict preferences in new sites, and estimate overall prey consumed in new sites.
... While potential predators of olive baboons (e.g. leopards, lions, and spotted hyenas Crocuta crocuta [57][58][59][60]) may also frequently use unpaved roads in protected areas, these large carnivores are primarily nocturnal in LMNP [61]. As we carried out the line distance surveys between 8 am and 6 pm, olive baboons likely perceived moving along roads during these times not only as convenient but also as relatively safe. ...
Estimating population density and population dynamics is essential for understanding primate ecology and relies on robust methods. While distance sampling theory provides a robust framework for estimating animal abundance, implementing a constrained, non-systematic transect design could bias density estimates. Here, we assessed potential bias associated with line distance sampling surveys along roads based on a case study with olive baboons (Papio anubis) in Lake Manyara National Park (Tanzania). This was achieved by comparing density estimates of olive baboons derived from road transect surveys with density estimates derived from estimating the maximum number of social groups (via sleeping site counts) and multiplying this metric with the estimated average size of social groups. From 2011 to 2019, we counted olive baboons along road transects, estimated survey-specific densities in a distance sampling framework, and assessed temporal population trends. Based on the fitted half-normal detection function, the mean density was 132.5 baboons km⁻² (95% CI: 110.4–159.2), however, detection models did not fit well due to heaping of sightings on and near the transects. Density estimates were associated with relatively wide confidence intervals that were mostly caused by encounter rate variance. Based on a generalized additive model, baboon densities were greater during the rainy seasons compared to the dry seasons but did not show marked annual trends. Compared to estimates derived from the alternative method (sleeping site survey), distance sampling along road transects overestimated the abundance of baboons more than threefold. Possibly, this overestimation was caused by the preferred use of roads by baboons. While being a frequently used technique (due to its relative ease of implementation compared to spatially randomized survey techniques), inferring population density of baboons (and possibly other species) based on road transects should be treated with caution. Beyond these methodological concerns and considering only the most conservative estimates, baboon densities in LMNP are among the highest across their geographic distribution range.
... Biomass of medium-sized prey has the strongest overall influence on spotted hyena biomass. Despite spotted hyenas being adaptable in the prey that they target (Hayward, 2006;Mills, 1990), this result is corroborated by known preferred prey weights of 56-182 kg (Hayward & Kerley, 2008), equivalent to small-to mediumsized prey in this study. ...
The spotted hyena (Crocuta crocuta Erxleben) and the lion (Panthera leo Linnaeus) are two of the most abundant and charismatic large mammalian carnivores in Africa and yet both are experiencing declining populations and significant pressures from environmental change. However, with few exceptions, most studies have focused on influences upon spotted hyena and lion populations within individual sites, rather than synthesizing data from multiple locations. This has impeded the identification of over-arching trends behind the changing biomass of these large predators. Using partial least squares regression models, influences upon population biomass were therefore investigated, focusing upon prey biomass, temperature, precipitation, and vegetation cover. Additionally, as both species are in competition with one other for food, the influence of competition and evidence of environmental partitioning were assessed. Our results indicate that spotted hyena biomass is more strongly influenced by environmental conditions than lion, with larger hyena populations in areas with warmer winters, cooler summers, less drought, and more semi-open vegetation cover. Competition was found to have a negligible influence upon spotted hyena and lion populations, and environmental partitioning is suggested, with spotted hyena population biomass greater in areas with more semi-open vegetation cover. Moreover, spotted hyena is most heavily influenced by the availability of medium-sized prey biomass, whereas lion is influenced more by large size prey biomass. Given the influences identified upon spotted hyena populations in particular, the results of this study could be used to highlight populations potentially at greatest risk of decline, such as in areas with warming summers and increasingly arid conditions.
Chimpanzees ( Pan troglodytes ) in Senegal may use nocturnality to mitigate hyperthermia risk in semi-arid environments but the degree of nocturnality for such chimpanzees also in sympatry with large carnivores remains uncertain. We compared diel activity among chimpanzees and their potential predators at Assirik in Niokolo-Koba National Park and contextualized these findings relative to other unit-groups in savanna landscapes. From 2015-2018, we generated a predator inventory using multi-modal methods and monitored the diel activity of chimpanzees and predators with camera traps [ camera trap (CT) days]. From 2015-2023, we also surveyed for evidence of predation during recce walks. Six potential nonhuman predators occur at Assirik, including lions ( Panthera leo ), leopards ( Panthera pardus ), spotted hyenas ( Crocuta crocuta ), African wild dogs ( Lycaon pictus ), Nile crocodiles ( Crocodylus niloticus ), and rock pythons ( Python sebae ). We documented one suspected case of a predator killing a chimpanzee. Nocturnality comprised 12.7% of CT events for chimpanzees and these events were more concentrated at twilight. Chimpanzees were more active during the day, predators were more active at night, and there was substantial temporal overlap among chimpanzees and potential predators during twilight intervals. Our findings support the hypothesis that savanna chimpanzees in Senegal are active at night in response to the extremely hot environment. We hypothesize that Assirik chimpanzees experience a tension between decreasing hyperthermia and increasing predation risk during nocturnality.
Energy flow through consumer-resource interactions is largely determined by body size. Allometric relationships govern the dynamics of populations by impacting rates of reproduction as well as alternative sources of mortality, which have differential impacts on smaller to larger organisms. Here we derive and investigate the timescales associated with four alternative sources of mortality for terrestrial mammals: mortality from starvation, mortality associated with aging, mortality from consumption by predators, and mortality introduced by anthropogenic subsidized harvest. The incorporation of these allometric relationships into a minimal consumer-resource model illuminates central constraints that may contribute to the structure of mammalian communities. Our framework reveals that while starvation largely impacts smaller-bodied species, the allometry of senescence is expected to be more difficult to observe. In contrast, external predation and subsidized harvest have greater impacts on the populations of larger-bodied species. Moreover, the inclusion of predation mortality reveals mass thresholds for mammalian herbivores, where dynamic instabilities may limit the feasibility of megafaunal populations. We show how these thresholds vary with alternative predator-prey mass relationships, which are not well understood within terrestrial systems. Finally, we use our framework to predict the harvest pressure required to induce mass-specific extinctions, which closely align with previous estimates of anthropogenic megafaunal exploitation in both paleontological and historical contexts. Together our results underscore the tenuous nature of megafaunal populations and how different sources of mortality may contribute to their ephemeral nature over evolutionary time.
Competition is known in biology through various mechanisms and species often compete for resources indirectly. We consider a system consisting of a predator, prey, which is not fully consumed and converts to a secondary resource (carcasses), and scavengers, which feed on the carcasses. Such an ecosystem is a simplified sketch of interactions in the savannah between lions and spotted hyenas, with lions killing herbivore prey, especially ungulates, whose leftovers serve as primary food for the hyenas. We perform an analytic and numerical study of the possible bifurcations which the model presents between states where one or more species are absent and the coexistence state.
Conservation planning has focused extensively on representation of species in protected areas (PAs), rather than achieving the persistence of these species in PAs. Consequently, most PAs do not support viable populations of many species represented within them, particularly large mammals, resulting in extinction debt risks. The challenge is therefore to identify opportunities to achieve viable populations and to develop and implement conservation plans to achieve this, e.g., expand and connect existing PAs. A recent meta-analysis provides a minimum viable population (MVP) estimate of 5 000 individuals for mammals. This value was used as the target for modelling conservation opportunities for the South African endemic Cape mountain zebra (Equus zebra zebra), blesbok (Damaliscus pygargus phillipsi), black wildebeest (Connochaetes gnou) and co-existing medium- and large-sized mammal species. This study focused on the region between the Camdeboo and Mountain Zebra national parks (ca. 736 000 ha). Within the envisaged Camdeboo Mountain Zebra (C-MZ) mega-reserve, a spreadsheet model was used to estimate potential herbivore population sizes, spatial requirement data were used to estimate potential omnivore and insectivore population sizes, and herbivore densities were used to estimate potential large carnivore population sizes. I show that 12 (the three endemic grazers and nine co-existing species) of the 28 modelled medium- and large-sized mammal species will potentially exceed 5 000 individuals, the MVP target, within the envisaged mega-reserve. The remaining 16 species show potential to reach substantial population sizes, however, will likely require ongoing metapopulation management. Using a conservation planning approach, I identify priority areas for the conservation of suitable habitat for Cape mountain zebra, blesbok, and black wildebeest within the envisaged mega-reserve, thus indirectly identifying priority areas for the conservation of these endemic species. I show that the potential incorporation of these priority areas within the envisaged mega reserve will contribute considerably to the conservation of all focal species. In addition, the potential incorporation of these priority areas will contribute to PA expansion and significantly improve connectivity between existing PAs within the envisaged C-MZ mega-reserve. The importance of adequately-sized PAs to support viable populations of focal species is thus evident, and will ensure their long-term survival and ability to persist by allowing population and evolutionary scale processes.
Once considered mere scavengers, it is now widely recognized that hunting is more important than scavenging in the feeding ecology of spotted hyenas (Crocuta crocuta). In this chapter, we outline the extraordinary morphological and behavioral adaptations possessed by these bone-cracking hyenas for efficient hunting and foraging within the context of their complex social organization. These social carnivores live in female-dominated societies structured by fission-fusion dynamics in which individuals hunt alone or in small groups to avoid feeding competition but join forces in large-scale cooperation with kin and non-kin group-mates to defend food from African lions (Panthera leo) and members of neighboring groups of hyenas. We discuss how social rank and age influence every aspect of their hunting behavior and consider the inevitable trade-offs faced regarding cooperative hunting of ephemeral prey. Finally, we evaluate what is known about the cognitive demands and conservation implications associated with the behavioral flexibility possessed by these efficient hunters.
To know the level of wildlife conservation in a protected area is fundamental for its rational management. In this study, abundance and distribution of mammalian fauna in the W Niger Transboundary Biosphere Reserve (RBTW Niger) was determined over five years. The determination of abundance consisted of identifying and counting the fauna along circuits in the protected area subdivided into four Counting Blocks, on the one hand, and settling 14 cameras trap, on the other hand. Sampling points were selected randomly. Thirty-one (31) species of mammalian fauna were identified during the surveys. The most observed species are hippotragues (15.22%),
oribis (12.57%), buffon's cobes (12.04%), baboon (10.99%), warthog and buffalo with respectively 8.75% and 7.62% of contacts. Carnivores in general and savannah lions in particular were less observed during this study, due to threats and pressures which they are subjected. In order to better understand large carnivores in the RBTW, a study was conducted to elucidate distribution of habitats of theirs preys (buffaloes and hippotragues) in general and for savannah lions in particular.
The method consisted of mapping habitats of these two preys using two climate models (CSIRO and CCMA) with the Maximum Entropy tool (Maxent), DIVA GIS and QGIS. Future distributions showed unfavorable habitats for the buffalo that fluctuates following a trend of +46.35 to +354.50% and very favorable habitats for the hippotrague with a trend of +129.39 to +166.31% compared to the distribution over the period 1950-2000. The RBTW can therefore provide suitable habitats for
hippotragues that lions must choose to ensure their survival. This will limit human-wildlife conflicts. Species involved in livestock predation and crops damage are carnivores groups with 59.52% of occurrence followed by herbivores with 26.98% and omnivores (13.49%). The analysis of wildlife abundance and distribution shows that RBTW Niger area is relatively well protected based on the diversity indices recorded in the Counting Blocks. The sustainable management of carnivores requires the monitoring and conservation of their habitats; it has been shown for instance that the conflict between human and wildlife on the outskirts of the Reserve is relatively limited.The use of new technologies such as trap cameras seems to provide results similar to those of survey; Distance Sampling's transect point method applied to trap camera data may be a breakthrough in wildlife monitoring.
Keywords: Abundance and distribution, wildlife conservation, Point transect, Habitat Modelling,
RBT W Niger.
Humans evolved in the dynamic landscapes of Africa under conditions of pronounced climatic, geological and environmental change during the past 7 million years. This book brings together detailed records of the paleontological and archaeological sites in Africa that provide the basic evidence for understanding the environments in which we evolved. Chapters cover specific sites, with comprehensive accounts of their geology, paleontology, paleobotany, and their ecological significance for our evolution. Other chapters provide important regional syntheses of past ecological conditions. This book is unique in merging a broad geographic scope (all of Africa) and deep time framework (the past 7 million years) in discussing the geological context and paleontological records of our evolution and that of organisms that evolved alongside our ancestors. It will offer important insights to anyone interested in human evolution, including researchers and graduate students in paleontology, archaeology, anthropology and geology.
Humans evolved in the dynamic landscapes of Africa under conditions of pronounced climatic, geological and environmental change during the past 7 million years. This book brings together detailed records of the paleontological and archaeological sites in Africa that provide the basic evidence for understanding the environments in which we evolved. Chapters cover specific sites, with comprehensive accounts of their geology, paleontology, paleobotany, and their ecological significance for our evolution. Other chapters provide important regional syntheses of past ecological conditions. This book is unique in merging a broad geographic scope (all of Africa) and deep time framework (the past 7 million years) in discussing the geological context and paleontological records of our evolution and that of organisms that evolved alongside our ancestors. It will offer important insights to anyone interested in human evolution, including researchers and graduate students in paleontology, archaeology, anthropology and geology.
Stable isotope palaeoecology of fossil mammals is a key research tool for understanding the environmental context of hominin evolution in the Plio-Pleistocene of Africa. Well studied mammal groups include bovids, suids, equids, proboscideans and primates, but to date there has been no in-depth study of modern and fossil carnivores. Here we produce an Africa-wide oxygen and carbon enamel isotope dataset for modern carnivores and compare it with fossil carnivore data sampled from the Plio-Pleistocene Omo Group of the Turkana Basin, Kenya. Comparison of modern carnivore carbon isotopes with satellite images of land cover indicates that carnivore δ¹³C is related to the proportion of woody cover in the local environment. Modern carnivore oxygen isotopes are strongly influenced by the δ¹⁸O of meteoric water, through drinking from standing water and through prey body fluids. Carbon isotope data from fossil carnivores shows close agreement with palaeovegetation reconstructions from δ¹³C of palaeosol carbonates from the same geological Members, and a similar long-term trend in δ¹³C values through time (4 Ma to 1 Ma), reflecting a gradual increase in the proportion of C4 grasses in the Turkana Basin. This increase in the δ¹³C of large carnivores is consistent with the evidence from other mammalian groups for an increase in the proportion of grazers compared to browsers and mixed feeders during this time interval. Two distinct trends within oxygen versus carbon isotope space indicates that the fossil carnivores lived during two distinct climatic regimes – one in which palaeo-lake Turkana was freshwater, and one in which the lake resembled its modern-day hyperalkaline state. These two climatic states most likely represent the end-members of precessionally-driven rainfall extremes over the Ethiopian Highlands. This indicates that each studied faunal assemblage from the Omo Group is a time- and climate-averaged palimpsest; this has significant implications for the interpretation of environmental signals and community palaeoecology derived from Turkana Basin fossil mammals, including early hominins.
The spotted hyena (Crocuta crocuta) is a large and unique terrestrial carnivore. It is a particularly fascinating species due to its distinct phenotypic traits, especially its complex social structure and scavenging lifestyle, with associated high dietary exposure to microbial pathogens. However, the underlying molecular mechanisms related to these phenotypes remain elusive. Here, we sequenced and assembled a high-quality long-read genome of the spotted hyena, with a contig N50 length of ∼13.75 Mbp. Based on comparative genomics, immunoglobulin family members (e.g., IGKV4-1) showed significant adaptive duplications in the spotted hyena and striped hyena. Furthermore, immune-related genes (e.g., CD8A, LAG3, and TLR3) experienced species-specific positive selection in the spotted hyena lineage. These results suggest that immune tolerance between the spotted hyena and closely related striped hyena has undergone adaptive divergence to cope with prolonged dietary exposure to microbial pathogens from scavenging. Furthermore, we provided the potential genetic insights underlying social complexity, hinting at social behavior and cognition. Specifically, the RECNE-associated genes (e.g., UGP2 and ACTR2) in the spotted hyena genome are involved in regulation of social communication. Taken together, our genomic analyses provide molecular insights into the scavenging lifestyle and societal complexity of spotted hyenas.
The literature on cheetah Acinonyx jubatus ecology is dominated by studies on the Serengeti Plains (SNP) in East Africa. Because of this and the cheetah's hunting strategy it is generally considered to be a predator that prefers open grassland plains. However, cheetahs also inhabit a range of bush, scrub and woodland habitats. A field study using direct observations of radio-collared individuals in the woodland savanna habitat of the Kruger National Park (KNP), South Africa, and a literature review of studies across savanna habitats examined cheetah predation, hunting behaviour and habitat use in relation to prey composition, cover availability and kleptoparasitism. The cheetah's main prey is medium-sized herbivores, with a bias towards male prey. The group size and sex of the hunting cheetah may influence the results of prey selection studies as male coalitions tend to take larger prey than females. Cheetahs initiated more hunts and had a higher success rate in the open woodland savanna of the KNP compared to other available habitats with thicker bush, and in other wooded savanna areas they also prefer more open habitat for hunting. Although they appear to have shorter chase distances in more wooded habitats, hunting success appears to be slightly higher in open grassland habitat. Woody vegetation appears to obstruct the cheetah's high-speed hunting strategy, thereby lowering hunting success. However, cheetahs use cover for stalking prey and open habitats with bordering woodlands, or patches of cover are considered preferred cheetah habitats. In these habitats, cheetahs can stalk closer to their prey using available cover, but also successfully pursue their prey into available open spaces. Across African savanna ecosystems, cheetahs appear to be kleptoparasitised less in more wooded habitats. Therefore they may also prefer these habitats because they provide greater concealment from kleptoparasites. Our study suggests that the cheetah is more adaptable to habitat variability than is often thought and is not only a successful hunter on open grassland plains.
Species composition of prey consumed by spotted hyaenas Crocuta crocuta in three divergent areas in southern Africa was determined by scat analyses. The larger abundant antelopes predominated in the diet and their occurrence in the diet was apparently directly related to the availability of the prey within the hyaena foraging areas. In Mkuzi' Game Reserve and the Namib Naukluft National Park, hyaenas coexist with only one other large predator. In Umfolozi Game Reserve, where hyaenas coexist with four other large predators, a greater variety of prey was taken. Scat weight in desert-dwelling hyaenas was twice that of those from more mesic areas, which may be an adaptation to restricted water intake. The incidence of domestic livestock in the diet was meagre, probably the result of collecting scats only from latrines within the game reserves. Die spesies-samestelling van prooi verbruik deur gevlekte hienas Crocuta crocuta in drie verskillende gebiede in Suidelike Afrika is bepaal deur faeces-analise. Die groter meer volop wildsbokke het die dieet gedomineer en die voorkoms in dieet toon 'n direkte verwantskap met beskik baarheid van prooi in hiena-jaggebiede. Hienas deel Mkuzi Wildtuin en die Namib Naukluft Nasionale Park met slegs een ander groot roofdier terwyl vier ander groot roofdiere Umfolozi Wildtuin met hienas deel, wat lei tot groter variasie in prooi-items. Faeces-massa van woestynlewende hienas ..,as twee keer meer as die van hienas in meer mesiese gebiede wat waarskynlik 'n aanpassing is vir beperkte water inname. Die voorkoms van plaasdiere in die dieet was baie laag, maar dit is heel waarskynlik as gevolg van faeces versameling slegs binne die wildtuine.
The feeding ecology of the spotted hyaena (Crocuta crocuta) was studied in dense mountain forest of Aberdare National Park, Kenya. 332 scats and 138 regurgitations were collected and prey hair and other contents identified. Bushbuck, suni and buffalo were the most important food titems, accounting for 59% of food occurrences in the samples. Availability of prey species was estimated from road counts. Hyaenas were the dominant predators in the ecosystem. The importance of hunting as opposed to scavenging by hyaenas is discussed.
The African wild dog Lycaon pictus is critically endangered, with only about 5,000 animals remaining in the wild(1). Across a range of habitats, there is a negative relationship between the densities of wild dogs and of the spotted hyaena Crocuta crocuta(2). It has been suggested that this is because hyaenas act as 'kleptoparasites' and steal food from dogs. We have now measured the daily energy expenditure of free-ranging dogs to model the impact of kleptoparasitism on energy balance, The daily energy expenditures of six dogs, measured by the doubly labelled water technique, averaged 15.3 megajoules per day. We estimated that the instantaneous cost of hunting was twenty-five times basal metabolic rate. As hunting is energetically costly, a small loss of food to kleptoparasites has a large impact on the amount of time that dogs must hunt to achieve energy balance. They normally hunt for around 3.5 hours per day but need to increase this to 12 hours if they lose 25% of their food. This would increase th
Habitat use of quokka (Setonix brachyurus) in the northern jarrah forest of Western Australia was determined by monitoring movements of 58 quokkas over 2 years in 5 local populations. Quokkas were largely restricted to Agonis swamps that occur patchily throughout the jarrah forest. Within swamps, they are habitat specialists, preferring early seral stages that have been burned within the previous 10 years. This preference derives from a combination of dietary requirements and refuge from predation. As swamps mature they become suboptimal, forcing quokkas to colonize new patches. Since the collapse of the metapopulation following the introduction of the European red fox (Vulpes vulpes) in the 1930s, quokkas have been forced to remain at a site because predation inhibits dispersal.
Valuable conservation research on the African wild dog (Lycaonpictus) has identified that its current endangerment is primarily due to human persecution, although habitat alteration, interference competition with other large predators, and disease also are factors. Numerous studies have thus determined what should be avoided to sustain an African wild dog population, yet in this study we identify what is needed to conserve a wild dog population by using Jacobs' index to determine its preferred prey species. Twenty-four assessments of wild dog prey preference were calculated from 18 studies involving 4,874 kills of 45 species from throughout its distributional range. Wild dogs prefer prey within a bimodal body mass range of 16-32 kg and 120-140 kg, which is abundant and less likely to cause injury when hunted. This bimodal range follows that of optimal wild dog pack sizes based on energetic costs and benefits. Greater kudu (Tragelaphus strepsiceros) and Thomson's gazelle (Gazella thomsonii) are killed by wild dogs wherever they coexist and are significantly preferred. Impala (Aepyceros melampus) and bushbuck (Tragelaphus scriptus) also are significantly preferred. Our results allow wildlife managers to more accurately assess the survival chances of reintroduced or small wild dog populations by determining if sufficient preferred prey are available. These techniques are applicable to all adequately studied large predators.
Over the last three decades the main prey species (wildebeest Connochaetes taurinus , zebra Equus burchelli , Thomson’s gazelle Gazella thomsoni , and Grant’s gazelle Gazella granti ) of spotted hyaenas Crocuta crocuta in the Ngorongoro Crater, Tanzania, substantially declined in numbers, whereas buffalo Syncerus caffer numbers increased strongly. This provided a ‘natural experiment’ to investigate how a generalist predator such as the spotted hyaena responds to long‐term changes in prey populations. Here we compare data on the feeding ecology of Crater hyaenas from the late 1990s (period II) with equivalent data from the late 1960s (period I).
Hyaenas showed strong hunting preferences for wildebeest calves and gazelle fawns during both periods I and II. Adult buffaloes, prey difficult to hunt for hyaenas, were rarely hunted in either period despite a large increase in buffalo numbers from period I to II.
Hyaenas exhibited a functional response to the changes in prey populations by killing more buffalo calves and adult wildebeest during period II, relatively easy prey categories for hyaenas to hunt.
The proportion of carcasses scavenged and acquired by kleptoparasitizing other predators increased from period I to II. This was facilitated by an increase in the relative number of lion kills available to hyaenas during the same period.
Hyaenas defended a similar proportion of their kills against kleptoparasitic attempts by lions during both periods. Since the ratio of kills hyaenas kleptoparasitized from lions in relation to kills lost to lions increased from period I to period II, hyaenas obtained more food resources from lions during period II than I.
The success of hyaenas kleptoparasitizing lions was influenced by the presence of male lions and the ability of hyaenas to recruit sufficient clan members to appropriate carcasses from lions.
This study is the first to demonstrate both a functional response of a predator to changes in the size of prey populations and an associated change in foraging behaviour of the predator.
The diet of African wild dogs, lions and spotted hyenas was studied in the woodland savannah of Faro National Park and surrounding hunting zones in northern Cameroon during the dry season. Faecal analysis revealed that the diet of the three large carnivores overlaps considerably. Frequencies of prey remains of large and medium-sized antelopes were highest, with Buffon's kob being by far the most common prey species. Carnivores seem to consume the most abundant prey species. No prey item of domestic livestock was found in the faeces. Faecal analysis has proved to be a useful method to reveal diet choice for large carnivores in northern Cameroon. Detailed information on the distribution and density of large carnivores and their prey is needed to give a better picture of the status of carnivores in northern Cameroon and to help resolve conflicts with livestock. Further investigations into the seasonal variation of predation and other factors structuring the large carnivore community in northern Cameroon are recommended.
Occurrence and diet of ten carnivorous predators (four falconiforms, four owls, and two foxes), and population levels of their mammalian prey, were monitored over 45 months at a semi-arid site in north-central Chile. Early in this period, small mammals irrupted and then declined markedly to a density 7% of that at peak. All four falconiforms (Buteo polyosoma, Falco sparverius, Geranoaetus melanoleucus, Parabuteo unicinctus) and one owl (Tyto alba) responded numerically to the decline in mammalian prey by virtually abandoning the study site. The three other owls (Athene cunicularia, Bubo virginianus, Glaucidium nanum) and the two foxes (Pseudalopex culpaeus and P. griseus) remained. With few exceptions, throughout the study predators maintained species-specific preferences among small mammal species regardless of the absolute and proportional abundance of these prev. At no time did the two prey species most responsible for the irruption (the rodents Phyllotis darwini and Akodon olivaceus) occur in predators' diets out of proportion to their estimated relative abundance in the field. Predators were clearly unable to prevent the irruption from occurring. Given the absence of a clear functional response to the most irruptive species, predators seemed unlikely to have been responsible for the observed crash. At present, however, predators may be prolonging the crash and delaying the return of small-mammal populations to typical densities.
1. Group hunting in social carnivores is generally thought to have evolved through natural selection for improved efficiency of prey capture, increased prey size apprehended or defence of the kill against intra- and interspecific kleptoparasitism. 2. We used a simple model to explore how variation in hunting group size or wild dogs in the Serengeti influences defence of kills against kleptoparasitism from spotted hyenas and the trade-off effects this has on intake rate per dog for a given prey size selected. 3. The analysis presented suggests that while kleptoparasitism substantially influences the amount of time a hunting group can access a kill, increases in access rime with increased hunting group size rarely fully compensate Cor the reduction in each dog's share of the carcass due to scramble competition among the dogs. 4. A profitability index, which includes limitations of the probability of capturing different sized prey, gut capacity, food depletion and access time, suggests that small hunting groups (1-2) would be particularly vulnerable to kleptoparasitism because they are usable to fully satiate themselves before spotted hyenas take over their kills. Intermediate-sized hunting groups may be most effective at meeting nutritional demands over a range of prey sizes. 5. While reasons for the recent extinction of the Serengeti wild dog population remain speculative, this paper contributes lo the debate by proposing that kleptoparasitism by spotted hyenas would have placed a major constraint on the ability of individual wild dog packs to recover from episodic disasters.
Predator-prey relationships amongst the larger mammals of the Kruger National Park
The diet, foraging behaviour, social organisation and social behaviour of the brown hyaena Hyaena brunnea and the spotted hyaena Crocuta crocuta, and the interactions between these two species in the southern Kalahari are discussed. The brown hyaena is a scavenger of a wide variety of vertebrate remains, supplementing its diet with wild fruits and insects and is well adapted to this arid region. The spotted hyaena is a hunter-scavenger of large and medium-sized mammals and is not found in such numbers in the southern Kalahari as is the brown hyaena. These differences in diet have led to the evolution of large differences in foraging behaviour, social organisation, denning behaviour and communication patterns in the two species; spotted hyaenas having a more highly developed social system and living in far larger territories than brown hyaenas. Spotted hyaenas are dominant to brown hyaenas, but because of their low density in the southern Kalahari, have little effect on the brown hyaena population there.
Five years of mortality data of impala Aepyceros melampus, blue wildebeest Connochaetes taurinus, buffalo Syncerus caffer and kudu Tragelaphus strepsiceros have been used to describe the minimum mortality profiles of the respective species in an open savanna system with the full compliment of predators. Predation is the principle cause of natural mortalities on the study site, Letaba Ranch, which is adjacent to the Kruger National Park. The principle cause of impala mortality are leopards Panthera pardus and wild dogs Lycaon pictus. Lion Panthera leo were major contributors to the mortality of wildebeest and buffalo. Anthrax Bacillus anthracis was the main cause of kudu mortality.
Species in the mammalian order Carnivora exhibit a huge diversity of life histories with body sizes spanning more than three orders of magnitude. Despite this diversity, most terrestrial carnivores can be classified as either feeding on invertebrates and small vertebrates or on large vertebrates. Small carnivores feed predominantly on invertebrates probably because they are a superabundant resource (sometimes 90% of animal biomass); however, intake rates of invertebrate feeders are low, about one tenth of those of vertebrate feeders. Although small carnivores can subsist on this diet because of low absolute energy requirements, invertebrate feeding appears to be unsustainable for larger carnivores. Here we show, by reviewing the most common live prey in carnivore diets, that there is a striking transition from feeding on small prey (less than half of predator mass) to large prey (near predator mass), occurring at predator masses of 21.5-25 kg. We test the hypothesis that this dichotomy is the consequence of mass-related energetic requirements and we determine the predicted maximum mass that an invertebrate diet can sustain. Using a simple energetic model and known invertebrate intake rates, we predict a maximum sustainable mass of 21.5 kg, which matches the point where predators shift from small to large prey.
Five methods for studying food habits of large African carnivores are evaluated. Fecal analysis is useful for a basic description of the diet, provided that an adequate sample of scats can be obtained. However, it is impossible to quantify the amount eaten and to determine the proportions of killed versus scavenged food. Tracking spoor in restricted habitats is useful for most species, except the brown hyena (Hyaena brunnea). Opportunistic, and for most species, radio-location observations are biased towards large prey animals, because small animals are eaten quickly, leaving no trace. However, the data can be used to study sex and age selection of adult prey. Direct observations provide accurate measurements of consumption rates, killing frequency, and prey selection, provided they can be carried out without disturbing predator or prey. Mixing data from incompatible techniques must be avoided.
This is an account of the life and habits of two species of Hyaena, the brown and the spotted, living in South Africa. It contains a considerable body of detailed knowledge, including painstaking nightwork, and it provides help in designing management strategies for species in need of protection. --- Part 1 Feeding ecology: food availability; brown hyaena diet; spotted hyaena diet; diets of the other large carnivores and ecological separation of the predators; the impact of predation on the prey populations; relations between hyaenas and other carnivorous animals. Part 2 Comparative foraging and feeding behaviour: activity patterns and resting sites; foraging group sizes; the use of senses during foraging; foraging for vegetable matter, birds' eggs and insects; brown hyaena hunting behaviour; spooted hyaena hunting behaviour; feeding behaviour. Part 3 Social structure and spatial organization: brown hyaena clans; spotted hyaena clans; nomadic males; land tenure system; factors affecting the sizes of social groups and territories. Part 4 Communication patterns and social interactions: visual and tactile communications and social interactions; vocalizations; scent marking. Part 5 The comparative denning behaviour and development of cubs: dens; development of cubs; sub-adults; functional considerations of denning behaviour in the Hyaenidae. Part 6 The individual in hyaena society: degrees of relatedness between clan members; brown hyaena society; spotted hyaena society; evolutionary trands in the social systems of the two species. Part 7 Relations between, and management considerations for, brown hyaenas and spotted hyaenas. Appendices: 1 - common and scientific names of species mentioned in the text; 2 - estimated numbers of some ungulates in the spotted hyaena study area; 3 - ageing criteria of ungulates based on eruption of teeth in bottom jaws and tooth wear; 4 - methods used to measure territory sizes; 5 - degrees of relatedness between clan members.
African wild dogs ( Lycaon pictus) are a seriously endangered species with only around 5000 individuals left. We examined the factors likely to be affecting the distribution and density of wild dogs living in the Kruger National Park, South Africa. We collected data, by traditional and satellite radio‐tracking, on diet selection, sizes of packs, dispersion and sizes of territories, and patterns of habitat selection for eight packs of wild dogs. In Kruger wild dogs specialize on impala ( Aepyceros melampus) and kudu ( Tragelaphus strepsiceros). They live in packs averaging about nine dogs and inhabit territories of 350–950 km ² . Neighboring territories overlap to some extent but packs meet very rarely. The dispersion pattern of food plays only a minor role in determining the density and distribution of dogs in the Kruger; dogs exist at their lowest density where their food is most abundant. This apparent paradox comes about because the dogs seem to avoid areas that, although prey‐rich, are heavily used by lions. This is a sensible strategy because lions are a major cause of dog mortality, accounting for some 39% of natural pup deaths and at least 43% of natural adult deaths. Wild dogs may be conserved most successfully in areas with moderate to low densities of lions and spotted hyenas, and management strategies should be careful not to promote these species to the detriment of wild dogs and other carnivores. The densities of lions and spotted hyenas should be important criteria when considering an area for the reintroduction of wild dogs and other similar sized carnivores.
Factores que Afectan la Densidad y Distribución de Perros Salvajes en el Parque Nacional Kruger
El perro salvaje africano ( Lycaon pictus) es una especie en serio peligro de extinción, con una población estimada de solamente 5000 individuos. Hemos examinado los factores que con mayor probabilidad afectan la distribución y densidad de los perros salvajes que habitan el Parque Nacional Kruger en Sudáfrica. Información sobre la selección de alimento, tamaño de grupo, dispersión y tamaño de territorio y patrones de selección de hábitat fué colectada usando métodos tradicionales y seguimiento con satélite para ocho grupos de perros salvajes en el parque. Esta especie se especializa en impala ( Aepyceros melampus) y kudu ( Tragelaphus strepsiceros). Vive en grupos de 9 individuos en promedio y habita un territorio de 350‐950 km ² . Territorios vecinos se sobreponen en cierta proporción, pero los encuentros entre diferentes grupos raras veces ocurren. El patrón de dispersión de la comida juega solo un pequeño papel en la determinación de la densidad en lugares donde la comida es mas abundante. Esta aparente paradoja es consecuencia de que los perros parecen evitar zonas que, a pesar de ser ricas en presas, también son muy usadas por leones (Panthera leo). Esta estrategia es sensata, pues esta especie es una de las mayores causas de mortalidad para los perros, alcanzando un 39% de mortalidad natural de los cachorros y al menos un 43% de la mortalidad natural de los adultos. Los perros salvajes podrían ser conservados con mas éxito en zonas con una densidad moderada o baja de leones y hienas manchadas (Crocuta crocuta). Las estrategias de conservación deben ser cuidadosamente estudiadas para evitar favorecer estas especies en detrimento de los perros salvajes y otras especies de carnívoros. La denisdad de leones y hienas manchadas deberá ser un criterio importante cuando se considere una zona para la reintroducción de los perros salvajes y otros carnívoros de similar tamaño.
The food habits of the spotted hyaena were studied during the dry season in relation to food availability in the Niokolo Koba National Park, Senegal. Frequency of occurrence of the different foods was quantified by means of scat analysis; species occurrence in the diet was compared with availability in the study area. Large and medium-sized mammals, such as the buffalo, the hartebeest and the warthog, were eaten more often than expected.
On a étudié les habitudes alimentaires de l'hyène tachetée pendant la saison sèche par rapport avec la disponibilité de la nourriture, dans le Parc National de Niokolo Koba, au Sénégal. On a quantifié la fréquence de la présence des différentes sources de nourriture en analysant les crottes; la présence des espèces a été comparée à leur disponibilité dans la région étudiée. Les mammifères de moyenne et de grande taille, comme le buffle, le bubale et le phacochère, sont mangés plus souvent qu'on ne s'y attendait.
Predator avoidance is likely to play a strong role in structuring species communities, even where actual mortality due to predation is low. In such systems, mortality may be low because predator avoidance is effective, and if the threat of predation is lifted then entire community structures may be altered. Where competition is intense, then competitor avoidance may have a similar impact on communities. Avoidance behaviors have been documented for a wide range of species, but this is the first attempt to document avoidance behavior within a large carnivore community. Audio playback techniques are used to examine the risk perceived by cheetahs from their two main competitors that are also their main predators, lions and hyenas. The results from these experiments show that cheetahs actively moved away from lion and hyena playback experiments, compared with dummy playbacks where no sound was played. Cheetahs showed no differences in their responses to playbacks dependent on their sex or reproductive status, suggesting they were responding principally to a competition rather than a predation threat. However, cheetahs were much less likely to hunt after competitor playbacks than after dummy playbacks, and this resulted in a lower kill rate after competitor playbacks, demonstrating that the perceived presence of competitors had a noticeable impact on the foraging rate of cheetahs. Furthermore, while cheetahs moved just as far following lion playbacks as after hyena playbacks, they spent significantly more time looking at the loudspeaker and were less likely to make a kill after lion playbacks, suggesting that cheetahs perceive lions to be a greater threat than hyenas. Key words: anti-predator behavior, carnivores, competition, foraging strategy, kleptoparasitism, playback experiments, predator avoidance, predation risk. (Behav Ecol 11:624-632 (2000))
1. Aspects of the influence of predation on the African buffalo Syncerus caffer were studied in Lake Manyara National Park, northern Tanzania, from 1981 to 1985. 2. During the study period, predation by lions was the most important cause of death in buffalo but occasionally major epidemics, such as rinderpest, can be by far the most important cause of death. 3. A comparison of Manyara with four other areas in Africa shows that the Manyara buffalo ran the same level of risk as those in these other areas, and that in periods in which epidemics or drought conditions are absent it was normal that ultimately some 90% of the large herbivores fell victim to predation. 4. Juvenile mortality in females was higher than in males but adult mortality in females was lower than in males. Adult bulls ran a much higher risk to be killed by lions (12.9% p.a.) than herd-living buffalo (4.4% p.a.); the latter category comprised cows and sub-adult bulls. 5. Risk of predation was highest near the ecotone between grasslands and structurally closed vegetations. Grasslands were as dangerous as woodlands and thickets, and only the mudflats were really safe. The chance to be killed fell steeply away from (potential) cover for lions. 6. Buffalo were more vigilant at night than during the day, peripheral animals more so than central ones. Adult bulls were more vigilant than other categories of buffalo. It is concluded that vigilance in buffalo was not restricted to predator detection. 7. The choice of feeding areas in buffalo was not dictated by predation risk alone.
The capture of prey by the African lion in Ngorongoro Crater, Tanzania, is examined. For analytic purposes the process is envisaged as a consecutive set of events: searching, stalking, attacking, and subduing prey.Initiation of prey capture activity by the lion arises from the interaction of degree of stimulation presented by the prey, timing of the activity cycle of the lion (diurnal or nocturnal), and its level of hunger. These factors yield thresholds for initiation which result in kills being directly related to prey density at high and low densities and inversely density dependent at intermediate prey densities.Success in stalking appears to depend primarily upon the failure of the prey to see the approaching lion until it is within its effective distance. Using data from actual stalks and contrived experiments, the probability is calculated of the lion being detected for a range of conditions.The attack involves a matching of the sprinting abilities of the predator and prey. To analyze this aspect in detail, the velocity curves of running are defined for the lion and four prey species.
ON the Serengeti Plains and in the Ngorongoro Crater (Tanzania), the spotted hyaena is the most numerous of the large carnivores, and a study of its feeding habits is being undertaken as part of the Serengeti Research Project since it was suspected that hyaenas influenced the numbers of wildebeest (Connochaetes taurinus), zebra (Equus quagga) and Thomson's gazelle (Gazella thomsonii). The study, which started in July 1964, aims at an evaluation of the total consumption of the hyaena-populations in the Serengeti and Ngorongoro Crater in relation to the abundance of prey, and at an investigation of the behavioural and ecological mechanisms involved in the interaction between the hyaena (as a scavenger or a predator) and its food species. This report deals (1) with some aspects of the distribution of hyaenas in the areas mentioned, and (2) with some of their feeding habits as revealed during the first 16 months of my investigation.
Lions Panthera
leo are generally thought to prey on medium to large ungulates. Knowledge of which species are actually preferred and which are avoided is lacking, however, as is an understanding of why such preference or avoidance may arise. An analysis of 32 studies over 48 different spatial locations or temporal periods throughout the distribution of the lion shows that it preferentially preys upon species within a weight range of 190–550 kg. The most preferred weight of lion prey is 350 kg. The mean mass of significantly preferred prey species is 290 kg and of all preferred species is 201 kg. Gemsbok, buffalo, wildebeest, giraffe and zebra are significantly preferred. Species outside the preferred weight range are generally avoided. Species within the preferred weight range that are not significantly preferred (such as roan, sable and eland) generally have features that reduce predation either morphologically (e.g. sable horns), ecologically (e.g. roan and sable occurring at low density), or behaviourally (e.g. the large herd size and increased vigilance of eland). Warthog are below the preferred weight range yet are taken in accordance with their availability and this is probably due to their sympatry with lion, their relatively slow evasion speed and their lower level of vigilance. Plots of prey preference against prey body mass follows a bell curve with a right skew that, we argue, is caused by collective hunting by lions of larger-bodied prey. Our methods can be used on all large predators and are likely to be useful in assessing competition in sympatric communities of predators, cooperative hunting and predicting predator diets. This will allow us to move beyond descriptive dietary studies to improve our predictive understanding of the mechanisms underlying predator–prey interactions.
The feeding behaviour of the Talek dan of spotted hyaenas in the Masai Mara National Reserve, Kenya, was monitored continuously for 7 years. Talek hyaenas adapted to large temporal variations in prey abundance by being opportunistic predators. During the first half of the year, the hyaenas fed on resident ungulates, and their diet consisted mainly of topi and Thomson's gazelles. Upon arrival of the migratory herds of wildebeest and zebra from the Serengeti, Talek hyaenas switched to feeding on the wildebeest which provided them with a superabundance of food for about 3 months. After the migratory animals returned to the Serengeti, Talek hyaenas experienced a period of reduced prey abundance due to the temporary dispersion of resident ungulates. At this time hyaenas hunted the few remaining wildebeest, and also increased their use of the remaining resident animals. Although Talek hyaenas were generally opportunistic in their feeding behaviour, they did exhibit clear dietary preference for larger pl ey species, particularly wildebeest. Finally, carrion comprised only 5% of the biomass consumed by Talek hyaenas, the lowest proportion of carrion in the diet of any Crocuta population studied to date.
The paper presents data on group size frequencies and mean group sizes in 11 species of ungulates and, for seven of them, more detailed information on the composition of groups in Tsavo East National Park, Kenya. In several species, mean group size was higher in a part of the study area with more open vegetation. No significant seasonal variations in mean group size were found, except in waterbuck which concentrated markedly in one location during the dry season. Minor local or seasonal variations in the proportions and/or sizes of different group types were evident in most species. In general, grouping patterns were consistent with the type of social organization found in other areas for the species concerned. Frequencies and types of interspecific associations, and the possible factors underlying their formation, are also discussed.
The abundance and spatial organization of spotted hyaenas (Crocuta crocuta) were studied in dense forest of Aberdare National Park, Kenya. Hyaenas were attracted to baits, using amplified tape recordings of hyaena calls. Using sightings of known hyaenas at bait sites and elsewhere population size was estimated with the Lincoln Index as 1·34 hyaena per km2, organized into two territorial clans. Mean clan size was 47. Mean group size of hyaenas moving together was 2·55. Home ranges were exclusive and averaged 32 km2.
On a étudié l'abondance et l'organisation spatiale de l'hyène tachetée (Crocuta crocuta) dans la forêt épaisse du Pare National d'Aberdare, Kenya. Les hyènes ont été attiréà des amorces avec des enregistrements amplifiés des appels d'hyène. En utilisant des vues d'hyènes connues à des sites d'amorces et ailleurs, on a estimé la taille de la population avec l'Index Lincoln à 1,34 hyène par km2, organisée en deux clans territoriaux. La taille moyenne de clan était 47. La taille moyenne de groupe d'hyènes qui se déplacent ensemble était 2,55. Les aires de répartition territoriaux étaient exclusifs et avaient un moyen de 32 km2.
The hunting technique of spotted hyaenas is described. In this study the hyaenas acquired three quarters of their food through hunting. The prey consisted primarily of antelope, warthogs and zebra foals, all of which weigh less than 150 kg. Most larger animals captured were suffering from previous injury.
Despite living in clans of up to 50 members hyaenas generally hunt in groups of 2 to 4, thus increasing the chance that some groups may locate the scattered herds which constitute most of their prey. Once a kill has been made nearby clan members converge to feed. Hyaenas are mainly nocturnal hunters, a habit relating to their avoiding high temperatures rather than their need for the cover of darkness, as hyaenas can hunt successfully in daylight on cool days. Their hunting technique is to approach the prey in fan formation, usually from downwind. An initial dash to scatter the herd is followed by visual selection of any vulnerable animal. The chase continues only if potential prey is detected, thus capture rates per hunt are low. Yet due to the persistence of the hyaenas many animals are taken. Preliminary evidence indicates that hyaena predation is most likely to affect both the resident populations of impala and, through predation of foals, the migratory population of zebras.
On décrit la technique de chasse de la hyène tachetée. Lors de cette etude, les hyénes ont obtenu trois quarts de leur nourriture en chassant. Les proies consistaient essentiellement en antilopes, phacochéres et jeunes zébres, pesant tous moins de 150 kgs. Les animaux plus grands qui étaient capturés souffraient d'un handicap antérieur.
Malgré qu'elles vivent en clans de plus de 50 individus, les hyénes chassent généralement en groupe de 2 a 4, augmentant ainsi la chance que certains groupes puissent localiser les troupeaux épars qui constituent la majorité de leurs proies. Dès qu'un animal a été tué, les membres du clan qui sont a proximite convergent pour manger. Les hyénes sont surtout des chasseurs nocturnes, habitude a relier plus a la crainte des hautes températures qu'au besoin d'être protegees par l'obscurité puisque les hyénes peuvent chasser avec succes en plein jour quand il fait frais. Leur technique de chasse consiste a approcher leur proie, sépartées en éventail, habituellement contre le vent. Une premére attaque, destinée a faire éclater le troupeau, est suivie par la sélection visuelle d'un animal vulnérable. La chasse ne continue que si elles en ont trouvé un; les taux de capture par assaut sont donc bas. Mais vu la persévérance des hyenes, beaucoup d'animaux sont pris. Une premiére évidence montre que la prédation par les hyénes affecte surtout les populations locales d'impalas et, par la prédation des jeunes, les populations nomades de zébres.
The African wild dog Lycaon pictus has declined dramatically over the past 30 years. Formerly distributed throughout sub-Saharan Africa, today c. 5000 wild dogs remain in total, mostly in southern and eastern Africa. Wild dogs’ decline reflects the expansion of human populations and the associated fragmentation of habitat available to wildlife. Because wild dogs live at very low densities, even ‘fragments’ covering thousands of square kilometres may not support viable populations. Furthermore, packs often range beyond the borders of reserves, so even nominally protected populations are often subject to persecution, road accidents, snaring and disease contracted from domestic dogs. Such edge effects mean that reserves smaller than c. 10,000 sq km will provide only incomplete protection. The highest priority for wild dog conservation, therefore, is to maintain and promote the contiguity of areas available to wildlife. Establishing cross-border parks and buffer zones, and encouraging game ranching on reserve borders, will all be beneficial. In smaller areas, protecting wild dogs requires that edge effects be mitigated by: (i) working with local farmers to limit persecution; (ii) controlling snaring; (iii) routing roads carrying high-speed traffic away from wildlife areas; and (iv) minimizing contact between wildlife and domestic dogs. Most of these measures will also benefit other wildlife.
This study examined the perceptual features of leopards (Panthera pardus) used as recognition cues by bonnet macaques (Macaca radiata) at three sites in southern India. Two of these sites were protected deciduous forest areas, the Mudumalai Wildlife Sanctuary and the Kalakad-Mundanthurai Tiger Reserve. The third study site was a predator-rare urban setting, Bangalore city. Four models were presented for 10-s intervals: (1) upright spotted leopard in a stalking profile with its head facing the subjects; (2) the same spotted model presented upside down; (3) a dark-brown leopard model with the same configuration as the spotted model; and finally (4), the dark-brown model presented upside down. The upright models characterized the two leopard morphs found in the wild - spotted and dark melanic. Inverted presentations of the spotted model examined the effects of the same configuration in an incongruous position to document whether leopard spots were still recognized. Differences between models were examined using alarm calls and flight elicitation as indices of fear. The spotted upright model was found to be the most feared, followed by the spotted upside-down model, the dark upright model, and finally the dark upside-down model. Analysis of when individuals looked at troop members to assess risks revealed that the spotted upright and dark upside-down models engendered significantly lower frequencies of information seeking; the upright model typically engendered immediate flight and the dark upside-down model was generally ignored. The spotted upside-down model and the dark upright model engendered high rates of information seeking. These results appeared to reflect the importance of both spots and felid configuration in leopard recognition. Lower responsiveness to the dark upright model might reflect lack of experience with this rare melanic form.