Anthropogenic disturbance induces opposing population trends in spotted hyenas and African lions

Large carnivore populations are declining worldwide due to direct and indirect conflicts with humans. Protected areas are critical for conserving large carnivores, but increasing human-wildlife conflict, tourism, and human population growth near these sanctuaries may have negative effects on the carnivores within sanctuary borders. Our goals were to investigate how anthropogenic disturbance along the edge of the Masai Mara National Reserve, Kenya, influences the demography and space-use of two large carnivore species that engage in intense interspecific competition. Here we document, in one disturbed region of the Reserve, a rapid increase in the population size of one large predator, the spotted hyena (Crocuta crocuta), but a striking concurrent decline in numbers of another, the African lion (Panthera leo). Anthropogenic disturbances negatively affected lion populations, and decreasing lion numbers appear to have a positive effect on hyena populations, indicated here by an increase in juvenile survivorship. We also saw an increase in the number of livestock consumed by hyenas. Our results suggest human population growth and indirect effects of human activity along Reserve boundaries may be effecting a trophic cascade inside the Reserve itself. These results indicate both top-down and bottom-up processes are causing a shift in the carnivore community, and a major disruption of guild structure, inside the boundaries of one of the most spectacular protected areas in Africa.     

Competitive interactions between spotted hyenas and lions in the Etosha National Park, Namibia

Spotted hyenas are successful hunters, but they also scavenge. Their main food competitors are lions. In the Etosha National Park, Namibia hyenas are unable to prevent kleptoparasitism by lions and fail to acquire kills from lions. The reasons are the small ratio of hyenas to female and subadult lions at kills and the presence of adult male lions. Because of the hyenas’ small clan sizes and large territories they seem to be unable to recruit sufficient clan members to take over lion kills or deter lions from their own kills. In Etosha, 71% of hyena mortality was due to lions; four cubs and one adult female hyena were killed by male lions during a 1‐year study. Hyenas have evolved adaptations against lions and initiate aggressive interactions with lions without the immediate availability of food, which is termed mobbing behaviour. Etosha hyenas initiated mobbing attempts when lions were near the hyena's communal den. Possibly, Etosha hyenas mobbed lions to distract lions from the hyenas’ den and their cubs and to warn their dependent offspring to hide from lions.     

Early life social and ecological determinants of global DNA methylation in wild spotted hyenas

Environmental factors early in life can have lasting influence on the development and phenotypes of animals, but the underlying molecular modifications remain poorly understood. We examined cross‐sectional associations among early life socioecological factors and global DNA methylation in 293 wild spotted hyenas (Crocuta crocuta) in the Masai Mara National Reserve, Kenya, grouped according to three age classes (cub, subadult and adult). Explanatory variables of interest included annual maternal rank based on outcomes of dyadic agonistic interactions, litter size, wild ungulate prey density and anthropogenic disturbance in the year each hyena was born based on counts of illegal livestock in the Reserve. The dependent variable of interest was global DNA methylation, assessed via the LUminometric Methylation Assay, which provides a percentage methylation value calculated at CCGG sites across the genome. Among cubs, we observed approximately 2.75% higher CCGG methylation in offspring born to high‐ than low‐ranking mothers. Among cubs and subadults, higher anthropogenic disturbance corresponded with greater %CCGG methylation. In both cubs and adults, we found an inverse association between prey density measured before a hyena was 3 months old and %CCGG methylation. Our results suggest that maternal rank, anthropogenic disturbance and prey availability early in life are associated with later life global DNA methylation. Future studies are required to understand the extent to which these DNA methylation patterns relate to adult phenotypes and fitness outcomes.     

Home range occupation and habitat use of sable antelope in the Okavango Delta region of northern Botswana

Animals selectively utilize their environments within a hierarchical framework. Our study addressed how the home ranges of sable antelope selectively incorporated the landscape and habitat types available to them. It was conducted in a region of northern Botswana where the sable population was expected to be thriving, in contrast to their threatened status in the wild in South Africa. The movements and habitat use of three neighbouring sable herds were recorded by global positioning system (GPS) telemetry during parts of the seasonal cycle in a region adjoining the seasonally flooded Okavango Delta. Total home range extents covered by these herds were larger than those found for sable in other areas, and local population densities, taking into account the herd sizes, were accordingly lower than in these other areas. Access to surface water appeared to be the main limitation on seasonal home range occupation. Almost all of the local vegetation types were utilized, but sable herds generally favoured dryland grassland during the dry season and floodplain grassland during the wet season, contrary to what we had expected. Hence, it appeared that local home range occupation and habitat use by the three sable antelope herds could be influenced more by interactions with potential competitors and predators than by intrinsic habitat suitability.     

Effects of an open refuse pit on space use patterns of spotted hyenas

Although the use of anthropogenic food sources by carnivores is well documented, few studies have investigated the potential for these sites to influence the movements of livestock predators. We capitalized on a natural experiment in which a refuse pit, utilized by a group of intensively monitored spotted hyenas ( Crocuta crocuta ) in the Masai Mara National Reserve, Kenya, was closed midway through monitoring efforts. We compared hyena space use patterns before and after closure, while controlling for other variables, to investigate the influence of the pit on home range size, core area size and location, and the proximity of hyenas to dens and the refuse pit itself. We also investigated the influence of prey abundance and the social ranks of individual hyenas on variation in frequency of pit utilization. We found home range size to increase and core area size to decrease following pit closure. Space use was clearly influenced by the pit, as the group core area included the refuse site only while it was in operation, and hyenas were found closer to the pit before than after its closure. The most common rank group observed feeding at the pit were low-ranking females, and regular pit users were more likely to be found near the pit during times of relative prey scarcity. We discuss the potential of human refuse at pastoral villages to increase livestock losses to hyenas.     

Fine‐scale ecological and anthropogenic variables predict the habitat use and detectability of sloth bears in the Churia habitat of east Nepal

Once widespread throughout the tropical forests of the Indian Subcontinent, the sloth bears have suffered a rapid range collapse and local extirpations in the recent decades. A significant portion of their current distribution range is situated outside of the protected areas (PAs). These unprotected sloth bear populations are under tremendous human pressures, but little is known about the patterns and determinants of their occurrence in most of these regions. The situation is more prevalent in Nepal where virtually no systematic information is available for sloth bears living outside of the PAs. We undertook a spatially replicated sign survey‐based single‐season occupancy study intending to overcome this information gap for the sloth bear populations residing in the Trijuga forest of southeast Nepal. Sloth bear sign detection histories and field‐based covariates data were collected between 2 October and 3 December 2020 at the 74 randomly chosen 4‐km² grid cells. From our results, the model‐averaged site use probability (ψ ± SE) was estimated to be 0.432 ± 0.039, which is a 13% increase from the naïve estimate (0.297) not accounting for imperfect detections of sloth bear signs. The presence of termite mound and the distance to the nearest water source were the most important variables affecting the habitat use probability of sloth bears. The average site‐level detectability (p ± SE) of sloth bear signs was estimated to be 0.195 ± 0.003 and was significantly determined by the index of human disturbances. We recommend considering the importance of fine‐scale ecological and anthropogenic factors in predicting the sloth bear‐habitat relationships across their range in the Churia habitat of Nepal, and more specifically in the unprotected areas.     

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

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

Coprologic survey of parasites of spotted hyenas (Crocuta crocuta) in the Masai Mara National Reserve,Kenya

Seventy fecal samples from spotted hyenas (Crocuta crocuta) in the Masai Mara National Reserve, Kenya were examined for parasite eggs and oocysts using sugar flotation. A total of nine parasite genera were identified, and all samples were positive for at least one parasite species. Most individuals were infected with Ancylostoma sp. and Spirometra sp., and these species had the highest median intensity of infection. Other parasites identified include Isospora sp., Taeniidae, Spirurida, Toxocara sp., Mesocestoides sp., Dipylidium sp., and Trichuris sp.     

生境变化对集合种群系统生态效应的影响

通过大量的数值模拟发现 :生境恢复或扩展将导致集合种群的强弱序由自然数的顺序规律演变为奇数种群强 -偶数种群弱 ,同时集合种群里的最优秀种群将迅速扩张、发展为更为强大的最优势种。而当生境遭受到破坏 (毁坏 ) ,集合种群里的最优秀种群将迅速地伦为最弱者。如果栖息地的毁坏率大于集合种群优势种对栖息地的占有率 ,不仅集合种群里的优势种群将不可避免地灭绝 ,伴随最优秀种群走向灭绝的种群依次还有第二、第三、第四强等的种群。同时 ,将导致集合种群的强弱序由自然数的顺序规律演变为偶数种群强 -奇数种群弱。     

Fecal glucocorticoids reflect socio-ecological and anthropogenic stressors in the lives of wild spotted hyenas

Our aim was to identify natural and anthropogenic influences on the stress physiology of large African carnivores, using wild spotted hyenas (Crocuta crocuta) as model animals. With both longitudinal data from a single social group, and cross-sectional data from multiple groups, we used fecal glucocorticoids (fGC) to examine potential stressors among spotted hyenas. Longitudinal data from adult members of a group living on the edge of the Masai Mara National Reserve, Kenya, revealed that fGC concentrations were elevated during two periods of social upheaval among adults, especially among younger females; however, prey availability, rainfall, and presence of lions did not influence fGC concentrations among hyenas. Our results suggested that anthropogenic disturbance in the form of pastoralist activity, but not tourism, influenced fGC concentrations among adult male hyenas; rising concentrations of fGC among males over 12 years were significantly correlated with increasing human population density along the edge of the group's home range. As hyenas from this social group were frequently exposed to anthropogenic disturbance, we compared fGC concentrations among these hyenas with those obtained concurrently from hyenas living in three other groups undisturbed by pastoralist activity. We found that fGC concentrations from the undisturbed groups were significantly lower than those in the disturbed group, and we were able to rule out tourism and ecological stressors as sources of variation in fGC among the populations. Thus it appears that both social instability and anthropogenic disturbance, but not the ecological variables examined, elevate fGC concentrations and represent stressors for wild spotted hyenas. Further work will be necessary to determine whether interpopulation variation in stress physiology predicts population decline in groups exposed to intensive anthropogenic disturbance.     

Sex differences in the play behavior of immature spotted hyenas, Crocuta crocuta

Social, locomotor, and object play were studied in a colony of five male and five female peer-reared spotted hyenas during 12 1-hr tests while the animals were 13-19 months of age. Animals were tested in both same-sex and mixed-sex groups and were stimulated to play by the introduction of fresh straw and sawdust bedding. Each test was videotaped and the frequency of each type of play was determined by a time sampling procedure. Females played more frequently than males, however, the category of play which was elevated depended upon the social context during testing. In same-sex tests the frequency of vigorous social play displayed by females markedly exceeded that by males, but no comparable sex difference appeared in mixed-sex tests. Females engaged in locomotor play more frequently than males in mixed-sex tests, but no comparable sex difference appeared in same-sex tests. No sex difference in object play was observed. Two male and two female hyenas were gonadectomized prior to the initiation of the present sequence of tests. The results suggest that gonadectomy during the prepubertal period does not affect the frequency of play behavior. However, the small sample sizes preclude any conclusive determination of the effects of these gonadectomies on play.     

Land use,rangeland degradation and ecological changes in the southern Kalahari,Botswana

Dual‐scale analyses assessing farm‐scale patterns of ecological change and landscape‐scale patterns of change in vegetation cover and animal distribution are presented from ecological transect studies away from waterpoints, regional remotely sensed analysis of vegetation cover and animal numbers across the southern Kalahari, Botswana. Bush encroachment is prevalent in semi‐arid sites where Acacia mellifera Benth. is widespread in communal areas and private ranches, showing that land tenure changes over the last 40 years have not avoided rangeland degradation. Herbaceous cover is dominated in intensively grazed areas by the annual grass Schmidtia kalahariensis Stent and in moderately grazed areas by the perennial grass Eragrostis lehmanniana Nees. Nutritious perennial grass species including Eragrostis pallens Hack. Ex Schinz remain prevalent in Wildlife Management Areas. Other ecological changes include the invasion of the exotic Prosopis glandulosa Torr. and dense stands of Rhigozum trichotomum Kuntze. in the arid southwest. Regional patterns of wildlife species show that the expansion of cattleposts and fenced ranches has led to large areas of low wildlife conservation value even in areas where cattle production is not practiced. Findings show the need for integrated landscape‐scale planning of land use if the ecological value and biodiversity of the southern Kalahari is to be retained.     

北部湾印太江豚密度估算及适宜栖息地预测

目前我国印太江豚 (Neophocaena phocaenoides) 的基础信息十分有限,2018—2020年对北部湾离岸较近的广西壮族自治区合浦儒艮国家级自然保护区及离岸较远的涠洲岛水域进行了截线抽样法船只考察。应用DISTANCE估算出儒艮国家级自然保护区印太江豚密度约为每平方千米0.273 (0.133 ~ 0.561) 头,数量约88头 (95%CI：43 ~ 181);涠洲岛水域印太江豚密度约为每平方千米0.100 (0.048 ~ 0.210) 头,数量约137头 (95%CI：65 ~ 286)。利用最大熵模型 (MaxEnt) 分析北部湾水域印太江豚的适宜栖息地分布及面积。MaxEnt模型训练集和测试集AUC值分别为0.980和0.927,模型拟合度较好,结果表明离岸距离和海洋深度是影响印太江豚栖息地分布的主要因子,印太江豚的适宜栖息地在北部湾东北部近岸区和海南西侧,适宜度超过0.5的栖息地面积达14 630.62 km²。2018—2020年印太江豚适宜栖息地面积较2013年大幅度下降,0.5适宜度的栖息地面积减少了40%。参考印太江豚2个群体的平均密度和北部湾适宜栖息地面积,粗略估计北部湾水域印太江豚约1 463 ~ 3 994头。由此可见,北部湾水域内印太江豚密度比较高,应引起足够的重视。建议首先进行区域性海洋环境的整体保护,重点关注鱼类资源的恢复及对人类活动尤其渔船的管控,并尽快建立新的自然保护区。     

Mountain lions on the prairie: habitat selection by recolonizing mountain lions at the edge of their range

Mountain lions (Puma concolor) have historically experienced large‐scale range contractions, but are beginning to recolonize portions of their former range. To reach potential suitable habitats in eastern North America, mountain lions need to move across the grassland and agriculture‐dominated habitats of the Great Plains, which are different from the forested areas associated with mountain lions in western North America. To inform restoration planning in this area, it is important to understand differences in mountain lion habitat selection in this “nontraditional” grassland habitat. We tracked GPS‐collared mountain lions in the Northern Great Plains of Montana and identified movement states (localized or exploratory) using behavioral change point analysis and net‐squared displacement. We compared habitat selection between the different states using step‐selection functions that included several environmental covariates. Similar to elsewhere throughout their range, across both movement states, mountain lions tended to select forested environments that were farther from human development. In contrast to more traditionally occupied mountainous regions, mountain lions in the Great Plains selected areas of lower elevations. They selected areas both near and far from water, but avoided riparian areas and selected more rugged environments when in exploratory movement states. This suggests that mountain lions in the Northern Great Plains are utilizing river corridors, particularly those with rough or broken topography during exploratory phases. To enhance future recolonization and connectivity of mountain lions to the east of our study area, we encourage managers to maintain and restore forest fragments along river corridors in the Great Plains.     

Androgens and the role of female "hyperaggressiveness" in spotted hyenas (Crocuta crocuta)

It has been suggested that female dominance in spotted hyenas evolved due to selection for "hyperaggressive" androgenized females. According to this view, virilized external genitalia of female hyenas developed as a byproduct of selection for "androgen-facilitated social dominance." The evidence that female hyenas have higher androgen levels than males or other female mammals is inconclusive. We compared concentrations of testosterone (T), 5alpha-dihydrotestosterone (DHT), and androstenedione (AE) from a population in the Serengeti. Females had significantly lower T and DHT levels than predispersal and postdispersal males. AE levels did not significantly differ between females and postdispersal males, but were significantly lower in predispersal males. Our results, and those from studies that have demonstrated that male and female hyenas experience similar levels of maternal androgens during fetal development, provide little support for a theory of selection for female dominance and hyperaggressiveness through enhanced secretion of androgens. Our data are consistent with an alternative view that female virilization is a byproduct of selection for precocial aggressive cubs of both sexes. According to this view, high investment in lactation favored selection for accelerated fetal development, high neonatal aggression, and facultative siblicide.     

Resource use and the nutritional status of sable antelope in the Okavango Delta region of northern Botswana

The resource‐use patterns and nutritional status of sable antelope herds were investigated in the Okavango Delta region of northern Botswana for comparison with those documented for declining sable antelope populations elsewhere in southern Africa. GPS collars recorded the relative use of floodplain, upland and wooded grassland habitats by the sable herds while VHF beacons facilitated locating the herds for direct observations on feeding. Surprisingly, the sable herds made greatest use of upland grasslands, rather than the floodplain grasslands exposed after floodwater had receded, during the dry season months. In the upland grasslands, they exploited tall, fibrous grass species that retained quite high levels of greenness through the dry season. This ability, together with partial use of the floodplain and some browsing on new leaves and flowers, helped maintain dietary nitrogen and phosphorus levels, as indicated by faecal nutrient levels, above maintenance thresholds through the dry season. Hence, the sable herds in the study region did not seem to be limited nutritionally under the rainfall and flooding conditions prevailing during the study.