Intraspecific interactions in a high‐density leopard population

Although less studied than interspecific interactions, interactions among members of the same species can influence space use and temporal activity. Using techniques commonly applied to the analysis of interspecific interactions—multispecies occupancy modeling and the analysis of temporal activity patterns—we studied intraspecific interactions within a high‐density population of Persian leopards (Panthera pardus saxicolor) in Tandoureh National Park, northeastern Iran. Using camera‐trap data, we investigated spatiotemporal interactions between male leopards, lone female leopards, and families (cubs/females with cubs). While we hypothesized that male and female leopards would display different temporal activity patterns, we did not predict spatial avoidance between these groups. We also predicted that leopard families would exhibit spatiotemporal avoidance from male leopards due to the risk of infanticide. Contrary to our expectations, we did not find any evidence for spatial or temporal avoidance between leopard families and adult male leopards. Male and lone female leopards exhibited positive pairwise co‐occurrence, consistent with reports of high overlap between male and female leopard home ranges. While a high level of overlap in temporal activity patterns was found between males/lone females and males/families, there was evidence for variation in the proportion of time each leopard group was active in particular periods of the diel cycle. Male leopards showed cathemeral activity, while lone females and families were more active during daylight hours. The application of these techniques to interactions within a species has improved understanding of the ecology and behavior of this endangered solitary carnivore.     

Movement,activity and hunting behaviour of leopards in the Cederberg mountains,South Africa

Few data are available on the behaviour of leopards in the absence of competing large predators and human impact, both of which are believed to influence leopard activity and movements. Remote camera traps and global positioning system (GPS) collars were used to quantify leopard activity in the Cederberg Mountains, seasonal and sexual differences in their movements, and determine whether nocturnal hunting success was related to lunar activity. Seventy‐seven per cent of camera‐trap photographs were at night, with a strong male bias (69%) in captures. Daily displacement using one location per day suggested that males moved significantly further than females. However, multiple locations (≥6 per day) showed no difference because males moved in a more linear fashion, but not further each day, than females. In the Cederberg Mountains, an open rocky habitat with low human impact and no competing predators, leopards were predominantly nocturnal, mainly hunting diurnal prey species. Hunting success was low: leopards travelled long distances between kills, with nocturnal hunting success higher on darker nights.     

Calibrating a camera trap–based biased mark–recapture sampling design to survey the leopard population in the N'wanetsi concession,Kruger National Park,South Africa

Estimating large carnivore abundance can be challenging. A biased leopard (Panthera pardus) population survey was conducted in the N'wanetsi concession in the Kruger National Park (KNP), South Africa, using motion‐sensitive camera traps from April to August 2008. Survey effort included 88 trapping occasions and 586 trap days. The survey yielded 24 leopard photographs, comprising fourteen adults of eleven males and three females. The capture rate was determined to be 24.4 trap days per leopard. Estimates of population abundance stabilized at approximately 500 trap days. Precision of population estimates began to stabilize after 378 trap days. We estimated that there were nineteen leopards in an area of 150 km². Leopard density was estimated at 12.7 leopards per 100 km². We explore the possibility of employing the methods used in this study to survey the leopard population in the KNP and surrounding areas.     

The effect of competing carnivores on the feeding behaviour of leopards (Panthera pardus) in an African savanna

Knowledge of competition dynamics among Africa’s large carnivores is important for conservation. However, investigating carnivore behaviour in the field can be challenging especially for species that are difficult to access. Methods that enable remote collection of data provide a means of recording natural behaviour and are therefore useful for studying elusive species such as leopards (Panthera pardus). Camera traps and Global Positioning System (GPS) collars are powerful tools often used independently to study animal behaviour but where their data are combined, the interpretation of a species’ behaviours is improved. In this study we used data from baited camera trap stations to investigate the feeding habits of leopards at Malilangwe Wildlife Reserve, Zimbabwe. We investigated the influence of spotted hyenas, lions and other competing leopards on the feeding duration of leopards using Generalized Linear Mixed Effects Modelling. To test the influence of competing predators on resting distances from bait sites, eight leopards were fitted with GPS collars. Results showed that leopards spent the shortest time feeding on the baits in the presence of competing male leopards compared to other predators while lion presence caused animals to rest farthest from bait sites. Interaction analysis indicated that small‐bodied leopards spent significantly shorter durations feeding when spotted hyenas were present. Our findings demonstrate that competition from guild carnivores has negative impacts on the food intake of leopards, which may have implications for fitness and survival. This study provides a snapshot of the competition dynamics at bait sites which may give insight to ecosystem level interactions among large carnivores in savanna ecosystems.     

新疆托木尔峰国家级自然保护区雪豹的种群密度

雪豹(Panthera uncia)是一种仅公布于中亚高山地区的珍稀濒危大型猫科动物,被IUCN红皮书列为濒危物种,并被收录入CITES公约附录Ⅰ,在中国雪豹被列为国家一级重点保护动物(杨奇森和冯祚建,1998).     

Kill rates and food consumption of leopards in Bardia National Park, Nepal

This is the first study presenting data on kill rates and food consumption among Asian leopardsPanthera pardus Linnaeus, 1758. In Bardia National Park, Nepal, we found leopard kills by searching within areas with clusters of locations from radio collared leopards (2 males and 1 female with 2 cubs aged 4–9 months). We used two tracking schemes, 24-h intensive radio-tracking and daily monitoring, and we defined food consumption as the product of average prey live weight and proportion consumed. The three leopards consumed 89.2 kg of meat from five chitalAxis axis, one domestic dog and two birds during 19 days of 24-h intensive tracking, rendering an average (± SE) daily food intake of 4.7 ± 0.3 kgper capita. Twenty-five prey items (14 chital, one muntjacMuntiacus muntjak, four primates and six birds) were found during 180 days of daily monitoring of the female. All edible biomass was consumed in all kills, except for three chital, and the rate of kill consumption was positively related to the age of her cubs. The average daily food consumption of the female was 4.0 ± 0.3 kg/day, the kill rate (days/kill) including all prey categories was 5.6 ± 0.4 days, and the kill rate of ungulates was 10.6 ±0.7 days. Our food consumption estimates are higher than reported from arid African environments. We suggest that the high food consumption rate in our study is a consequence of a release from time-energy constraints due to high prey abundance.     

三江源国家级自然保护区内云塔村雪豹种群动态

种群监测可为物种研究和保护提供关键信息和依据。雪豹(Panthera uncia)作为亚洲高山生态系统的顶级捕食者和旗舰种, 一直是研究和保护的重点, 但其难以到达的栖息地、隐秘的行踪和广阔的家域使其监测工作开展难度较大, 雪豹种群动态研究较为匮乏。本研究在2013年10月至2019年1月期间, 使用当地社区维护的红外相机, 监测三江源国家级自然保护区通天河沿保护分区内青海省玉树州哈秀乡云塔村雪豹种群的密度和动态, 共识别出35只雪豹个体。基于数据质量较好的2015、2016、2017年连续3年的红外相机数据各年截取3个月数据, 使用空间标记-重捕模型估算种群数量和密度, 发现当地雪豹种群和成年个体密度基本维持稳定, 种群增长率为1.02, 但监测期间雪豹个体更替明显, 平均个体更替率为0.44, 并且围绕两片雪豹核心利用区域发生了领域取代。推测雪豹种群具有较多个体更替和领域取代是因为种群处在雪豹潜在扩散通道上, 或调查范围未覆盖完整种群。本研究是国内首次对雪豹进行较为长期的种群动态监测和分析, 研究结果体现了动态监测的重要性, 也显示出以当地社区为主体监测哺乳动物种群的可能性。     

泰国Khao Yai国家公园豹猫和云豹的生态和保护

于1997年10月到1999年10月在泰国KhaoYai国家公园对两种同域分布的物种（即豹猫和云豹）进行了研究。使用带诱饵的陷阱捕捉了这些动物,进行麻醉,确定了性别、年龄,并戴上了无线电项圈,对6只雌性豹猫和4只雄性豹猫进行了1—18个月的无线电追踪。豹猫在干旱季节的利用区域大于雨季,雄性利用的区域大于雌性。在夜间和晨昏时节,豹猫活动增加,但是并没有节律性活动。所有豹猫个体在旱季和雨季的活动都相似,但雄性个体的白天活动多于雌性。豹猫的行走距离有性别差异但是没有季节差异;其食物以鼠类为主。分别对1只雌性云豹和1只雄性云豹进行了17个月和7个月的无线电追踪,雌性个体的活动面积为39．4km^2,而雄性个体活动面积为42．2km^2,核心区都是2．9km2。云豹对半绿林的使用大于其它类型的植被,在晨昏和夜间的活动增加,其活动无节律。     

Modelling potential habitat suitability for critically endangered Arabian leopards (Panthera pardus nimr) across their historical range in Saudi Arabia

Camera trapping can detect and monitor rare species in landscapes spanning thousands of square kilometres but placement of cameras in areas where the animals most likely occur will increase detection success. This vital information is lacking for the critically endangered Arabian leopard (Panthera pardus nimr) that has undergone a 90% decline across its range in Saudi Arabia. We aimed to identify suitable Arabian leopard habitat and potential population capacity in Saudi Arabia using data from leopards living in ecologically analogous habitat in South Africa and Oman. We developed a resource selection function (RSF) from 14 leopards’ GPS data in the Cederberg, South Africa, and validated the model using three leopards in the Little Karoo, and two Arabian leopards in Oman. We then projected the model to the historical range of Arabian leopards in Saudi Arabia to estimate likely leopard locations and potential population sizes based on home range metrics. The RSF successfully discriminated between used and available locations (specificity = 96.7%) and had high predictive ability (Rho > 0.9). Leopards selectively used areas away from human settlements and roads, with high enhanced vegetation index, and intermediate slopes and elevations. Saudi Arabia could theoretically host 4 distinct populations totalling 162–362 Arabian leopard females, depending on home range size. Camera traps deployed in the south-western mountains of Saudi Arabia may be most likely to detect remnant populations of Arabian leopards. Further research is needed into the local abundance of prey species and human activity to ensure the persistence of suitable leopard ranges and inform conservation actions.     

Low leopard populations in protected areas of Maputaland: a consequence of poaching,habitat condition,abundance of prey,and a top predator

Identifying the primary causes affecting population densities and distribution of flagship species are necessary in developing sustainable management strategies for large carnivore conservation. We modeled drivers of spatial density of the common leopard (Panthera pardus) using a spatially explicit capture–recapture—Bayesian approach to understand their population dynamics in the Maputaland Conservation Unit, South Africa. We camera‐trapped leopards in four protected areas (PAs) of varying sizes and disturbance levels covering 198 camera stations. Ours is the first study to explore the effects of poaching level, abundance of prey species (small, medium, and large), competitors (lion Panthera leo and spotted hyenas Crocuta crocuta), and habitat on the spatial distribution of common leopard density. Twenty‐six male and 41 female leopards were individually identified and estimated leopard density ranged from 1.6 ± 0.62/100 km² (smallest PA—Ndumo) to 8.4 ± 1.03/100 km² (largest PA—western shores). Although dry forest thickets and plantation habitats largely represented the western shores, the plantation areas had extremely low leopard density compared to native forest. We found that leopard density increased in areas when low poaching levels/no poaching was recorded in dry forest thickets and with high abundance of medium‐sized prey, but decreased with increasing abundance of lion. Because local leopard populations are vulnerable to extinction, particularly in smaller PAs, the long‐term sustainability of leopard populations depend on developing appropriate management strategies that consider a combination of multiple factors to maintain their optimal habitats.     

四川东南地区云豹(Neofelis nebulosa)分布区变化及其成因

物种的分布变化是衡量生物多样性变化的一个重要层面,也是生物多样性变化研究领域备受关注的热点。根据对四川省东南地区6县14个乡镇的访问调查,分析了四川省东南部云豹（Neofelis nebulosa）分布区的变化趋势,并对近10年潜在有无云豹分布的区域在1988年和2017年的景观格局进行对比分析,探讨了四川东南地区云豹分布区变化的原因。结果显示：（1）40年前云豹曾在调查区域广泛分布,近40年来云豹分布范围呈现逐渐缩减的趋势,到近10年潜在有云豹分布的仅3个县3个乡镇;（2）天然阔叶林是云豹适宜的生境,潜在有云豹分布地区阔叶林在近30年里均是面积最大的景观类型,30年间增加了18.13%。潜在无云豹分布地区阔叶林在30年前占比为13.67%,30年间减少了39.29%,其主要景观类型在30年间由农业用地转换为竹林类型;（3）30年间,潜在有无云豹分布地区阔叶林破碎化程度均加剧,而潜在无云豹分布地区的破碎化程度较潜在有云豹分布地区严重;（4）潜在有无云豹分布区域居民野生动物保护意识均不强烈,在潜在无云豹分布区域的保护意识和意愿稍高;（5）本研究认为,早在40年前由于人为捕杀和土地开垦,就导致云豹种群在研究区域内衰退,近30年虽然农业用地大幅转换为竹林使森林总面积增加,但并未对云豹的保护起到积极作用,阔叶林的生境退变则使前期已经衰退的云豹种群分布区持续缩减,以致难觅踪迹。     

Weights of gaur (Bos gaurus) and banteng (Bos javanicus) killed by tigers in Thailand

The primary prey of tigers across much of South‐East Asia has been depleted, reducing the ability of already limited habitat to support tigers. To better understand the extent to which two of the largest prey species, gaur (Bos gaurus) and banteng (Bos javanicus), contribute to the tiger's diet, we estimated the average size of these species killed by tigers. This information is needed to more accurately calculate biomass of these species in the tiger's diet and to devise strategies to increase tiger carrying capacity where habitat is fragmented and limited in west‐central Thailand. We used temporally clumped locations of 24 satellite radio‐collared tigers to identify their kill sites and obtained mandibles from 82 gaur and 79 banteng. Kills were aged by teeth eruption sequence, sectioning the M1 molar and counting cementum annuli. Of all gaur killed, 45.2% were adults; of all banteng killed, 55.7% were adults. The average weight of banteng killed was 423.9 kg, which was similar to the 397.9 kg average weight for gaur. The mean weight of both prey species is 3.5–4.5 times greater than the predicted 1:1 preferred prey to predator ratio. In the absence of medium‐sized prey, killing these larger animals may be especially critical for female tigers provisioning nearly independent young when male offspring are already larger than the mother. This is the first study to present data on the average weights of gaur and banteng killed in South‐East Asia, and these results suggest that these are key prey species to target in tiger prey recovery efforts.     

Sampling bias in snow leopard population estimation studies

Accurate assessments of the status of threatened species and their conservation planning require reliable estimation of their global populations and robust monitoring of local population trends. We assessed the adequacy and suitability of studies in reliably estimating the global snow leopard (Panthera uncia) population. We compiled a dataset of all the peer-reviewed published literature on snow leopard population estimation. Metadata analysis showed estimates of snow leopard density to be a negative exponential function of area, suggesting that study areas have generally been too small for accurate density estimation, and sampling has often been biased towards the best habitats. Published studies are restricted to six of the 12 range countries, covering only 0.3–0.9% of the presumed global range of the species. Re-sampling of camera trap data from a relatively large study site (c.1684 km²) showed that small-sized study areas together with a bias towards good quality habitats in existing studies may have overestimated densities by up to five times. We conclude that current information is biased and inadequate for generating a reliable global population estimate of snow leopards. To develop a rigorous and useful baseline and to avoid pitfalls, there is an urgent need for (a) refinement of sampling and analytical protocols for population estimation of snow leopards (b) agreement and coordinated use of standardized sampling protocols amongst researchers and governments across the range, and (c) sampling larger and under-represented areas of the snow leopard's global range.     

Infanticide in male laboratory mice: Effects of social status,prior sexual experience,and basis for discrimination between related and unrelated young

Several aspects of infanticide, including effects of social status, prior sexual experience, and the basis for discrimination between related and unrelated young, were examined in Swiss-Webster laboratory mice (Mus musculus). Strange males introduced into the female's cage for 24 h on day 1 postpartum significantly reduced pup survival whereas the introduction of sires did not. Direct observations of infanticide were frequent, and the motor patterns used by males to kill pups are described. Males killed their own offspring when those young were in the nest of a strange female, whereas most males did not kill unrelated young in the nest of a familiar female. Thus, past association with the mother appears to be the single most important factor mediating male discrimination of young. Prior contact with a specific female's urine reduced a male's propensity for subsequently killing her young. The act of copulation itself also reduced infanticide. Infanticidal behaviour was correlated with the male's social status: most dominant males killed unrelated pups, whereas none of the subordinates engaged in infanticide. These results are discussed in terms of the population biology of this species.     

Leopard (Panthera pardus) occupancy in the Chure range of Nepal

Conservation of large carnivores such as leopards requires large and interconnected habitats. Despite the wide geographic range of the leopard globally, only 17% of their habitat is within protected areas. Leopards are widely distributed in Nepal, but their population status and occupancy are poorly understood. We carried out the sign‐based leopard occupancy survey across the entire Chure range (~19,000 km²) to understand the habitat occupancy along with the covariates affecting their occupancy. Leopard signs were obtained from in 70 out of 223 grids surveyed, with a naïve leopard occupancy of 0.31. The model‐averaged leopard occupancy was estimated to be 0.5732 (SE 0.0082) with a replication‐level detection probability of 0.2554 (SE 0.1142). The top model shows the additive effect of wild boar, ruggedness, presence of livestock, and human population density positively affecting the leopard occupancy. The detection probability of leopard was higher outside the protected areas, less in the high NDVI (normalized difference vegetation index) areas, and higher in the areas with livestock presence. The presence of wild boar was strong predictor of leopard occupancy followed by the presence of livestock, ruggedness, and human population density. Leopard occupancy was higher in west Chure (0.70 ± SE 0.047) having five protected areas compared with east Chure (0.46 ± SE 0.043) with no protected areas. Protected areas and prey species had positive influence on leopard occupancy in west Chure range. Similarly in the east Chure, the leopard occupancy increased with prey, NDVI, and terrain ruggedness. Enhanced law enforcement and mass awareness activities are necessary to reduce poaching/killing of wild ungulates and leopards in the Chure range to increase leopard occupancy. In addition, maintaining the sufficient natural prey base can contribute to minimize the livestock depredation and hence decrease the human–leopard conflict in the Chure range.     

Assessment of human–leopard conflict in Machiara National Park,Azad Jammu and Kashmir,Pakistan

Livestock depredation by leopards is one of the key conservation issues in landscapes with limited resources worldwide. Any attempts to mitigate human–leopard conflict and conserve the species in conflict should be based on an unequivocal understanding of the conflict patterns. A household survey was conducted with an intention to quantify livestock losses resulting from depredation by leopards in and around the Machiara National Park, Azad Jammu and Kashmir, Pakistan. A total of 301 livestock were killed between June 2007 and August 2008 by leopards. The extent of loss varied in different months, with maximum killing occurring in the month of May. Small-bodied livestock such as goats and sheep were more vulnerable than large-bodied cattle. Goats were killed in significantly higher numbers as compared to other livestock. Domestic animals were killed more frequently at night as compared to other times of the day. There was a significant effect of the nighttime protection measures on depredation. Most attacks happened in valleys and in areas far from the forests. Factors most closely associated with livestock depredation included decline of natural prey, herding practices, guarding especially during the nighttime, and repeated use of pastures where predators were known to be hunting actively. Attitudes of the local community towards leopards were largely negative, and further efforts should be made to improve support for carnivore conservation. We suggest widespread local community education and proactive human–leopard conflict management, particularly through adopting carnivore-friendly livestock protection measures.     

Clouded leopard (<Emphasis Type="Italic">Neofelis diardi</Emphasis>) predation on proboscis monkeys (<Emphasis Type="Italic">Nasalis larvatus</Emphasis>) in Sabah,Malaysia

In this study, we have reported two direct observations of individuals from a one-male group of proboscis monkeys (Nasalis larvatus) being killed by clouded leopards (Neofelis diardi) in the riverine forest along the Menanggul river, a tributary of the Kinabatangan river in Sabah, Malaysia. One of the two individuals was an infant female and the other was a juvenile female. Based on literature reviews and the observations reported here, we suggest that clouded leopard and crocodile might be significant potential predators of proboscis monkeys of any age or sex and that predation threats elicit the monkeys' anti-predator strategies. Moreover, the observations of the monkeys' behaviour when the group is attacked by a predator suggest that the adult males in one-male groups play an important role as protectors.     

Vigorous Dynamics Underlie a Stable Population of the Endangered Snow Leopard Panthera uncia in Tost Mountains,South Gobi,Mongolia

Population monitoring programmes and estimation of vital rates are key to understanding the mechanisms of population growth, decline or stability, and are important for effective conservation action. We report, for the first time, the population trends and vital rates of the endangered snow leopard based on camera trapping over four years in the Tost Mountains, South Gobi, Mongolia. We used robust design multi-season mark-recapture analysis to estimate the trends in abundance, sex ratio, survival probability and the probability of temporary emigration and immigration for adult and young snow leopards. The snow leopard population remained constant over most of the study period, with no apparent growth (λ = 1.08+−0.25). Comparison of model results with the “known population” of radio-collared snow leopards suggested high accuracy in our estimates. Although seemingly stable, vigorous underlying dynamics were evident in this population, with the adult sex ratio shifting from being male-biased to female-biased (1.67 to 0.38 males per female) during the study. Adult survival probability was 0.82 (SE+−0.08) and that of young was 0.83 (SE+−0.15) and 0.77 (SE +−0.2) respectively, before and after the age of 2 years. Young snow leopards showed a high probability of temporary emigration and immigration (0.6, SE +−0.19 and 0.68, SE +−0.32 before and after the age of 2 years) though not the adults (0.02 SE+−0.07). While the current female-bias in the population and the number of cubs born each year seemingly render the study population safe, the vigorous dynamics suggests that the situation can change quickly. The reduction in the proportion of male snow leopards may be indicative of continuing anthropogenic pressures. Our work reiterates the importance of monitoring both the abundance and population dynamics of species for effective conservation.     

Space–time clusters for early detection of grizzly bear predation

Accurate detection and classification of predation events is important to determine predation and consumption rates by predators. However, obtaining this information for large predators is constrained by the speed at which carcasses disappear and the cost of field data collection. To accurately detect predation events, researchers have used GPS collar technology combined with targeted site visits. However, kill sites are often investigated well after the predation event due to limited data retrieval options on GPS collars (VHF or UHF downloading) and to ensure crew safety when working with large predators. This can lead to missing information from small‐prey (including young ungulates) kill sites due to scavenging and general site deterioration (e.g., vegetation growth). We used a space–time permutation scan statistic (STPSS) clustering method (SaTScan) to detect predation events of grizzly bears (Ursus arctos) fitted with satellite transmitting GPS collars. We used generalized linear mixed models to verify predation events and the size of carcasses using spatiotemporal characteristics as predictors. STPSS uses a probability model to compare expected cluster size (space and time) with the observed size. We applied this method retrospectively to data from 2006 to 2007 to compare our method to random GPS site selection. In 2013–2014, we applied our detection method to visit sites one week after their occupation. Both datasets were collected in the same study area. Our approach detected 23 of 27 predation sites verified by visiting 464 random grizzly bear locations in 2006–2007, 187 of which were within space–time clusters and 277 outside. Predation site detection increased by 2.75 times (54 predation events of 335 visited clusters) using 2013–2014 data. Our GLMMs showed that cluster size and duration predicted predation events and carcass size with high sensitivity (0.72 and 0.94, respectively). Coupling GPS satellite technology with clusters using a program based on space–time probability models allows for prompt visits to predation sites. This enables accurate identification of the carcass size and increases fieldwork efficiency in predation studies.