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Prey preference of the tiger Panthera tigris
Abstract
Tigers Panthera tigris continue to decline despite the best efforts of the worldwide scientific and conservation communities. Prey depletion has been linked to this decline, but a clear definition of what constitutes preferred prey and preferred prey weight range does not exist. This is critical information if we are to assess tiger reintroduction potential, monitor unforeseen poaching of predators and prey, and successfully conserve the species. Here we reviewed the available literature on tiger diet and prey availability and calculated Jacobs's electivity index scores from 3187 kills or scats of 32 prey species. We found that wild boar and sambar deer are significantly preferred by tigers, with red deer and barasingha likely to be significantly preferred also with a larger sample size. Prey body mass was the only variable that related to tiger prey preference with species weighing between 60 and 250 kg preferred by tigers yielding a ratio of predator to preferred prey of 1:1, which is similar to other solitary felids. This information can be used to predict tiger diet, carrying capacity and movement patterns, as it has been for Africa's large predator guild, and has important implications for tiger conservation throughout its distribution.
... A global review study on tigers' food habit and prey selection pattern found that wild pig and sambar were largely consumed by tigers, with red deer and barasingha likely to be significantly preferred (Hayward et al., 2012). Prey body mass was one of the most important variables that related to tiger prey preference with species weighing between 60 and 250 kg found preferred by tigers yielding a ratio of predator to preferred prey of 1:1 (Hayward et al., 57 | P a g e 2012). ...
... Thereafter, the preliminary study on the reintroduced tigers in Sariska (Sankar et al., 2010) reported prey preference in the order of sambar> chital > nilgai> livestock (cattle and buffalo) > common langur. Hayward et al., (2012) reviewed the available worldwide literature on tiger diet based on prey availability and estimated Jacobs's electivity index scores from kill and scat data. The study found that wild pig (Sus scrofa) and sambar (Rusa unicolor) are preferred by tigers while red deer (Cervus elaphus) and barasingha (Cervus duvauceli) were much higher in term of preference. ...
... Since tiger is a specialized hunter (Sunquist and Sunquist, 1989), it can survive on a wide range of prey species (Hayward et al., 2012). Though they are considered habitat generalists (Sunquist, et al., 1999), tigers are still vulnerable to habitat loss and fragmentation since they always prefer a high biomass of large-sized prey (Karanth and Sunquist, 1995) and in the absence of wild prey populations their ranging pattern and resource selection gets really affected (Karanth, 1991). ...
In a forest ecosystem, large felids are generally the top predators in almost every food chain influencing the structure and dynamics of the subsequent descending trophic levels. Evaluation of last fifty years’ presence and absence status of large felids in Protected Areas (PAs) of Indian subcontinent revealed that local extinction was highest in the dry deciduous habitat. Tiger, among all large felids had already become locally extinct from 70% of semi-arid dry thorn and 35% of dry deciduous forest areas. Tigers in dry semi-arid forests with its global western most limit, survive in small isolated populations. The present study assessed the movements and ranging patterns, prey availability, prey utilization and resource selection of reintroduced tigers in Sariska Tiger Reserve from July 2008 to June 2012.
... Given DPKY's history, the interaction of environmental, prey and human factors on recent tiger distribution is likely complex. In tiger studies elsewhere in their range, prey emerges as the strongest, or among the strongest, predictors of tiger presence (Karanth et al., 2011;Harihar & Pandav, 2012;Ngoprasert et al., 2012;Barber-Meyer et al., 2013) with evidence suggesting an optimal tiger-prey body mass ratio of approximately 1:1 (~60-250 kg; Hayward et al., 2012). Conversely, studies also report strong negative associations between tiger presence and anthropogenic disturbance, such as roads (Kerley et al., 2002) and proximity to settlements (Sunarto et al., 2012), though these relationships may not be ubiquitous (Carter et al., 2012). ...
... In Western Thailand, studies have suggested wild boar (Steinmetz et al., 2013), gaur, sambar and banteng to be important prey species (Petdee, 2000;Prommakul, 2003). Wild boar and deer species, such as sambar, are suggested to be important prey species elsewhere in the tiger's range (Sunquist, Karanth, & Sunquist, 1999;Biswas & Sankar, 2002;Hayward, Jedrzejewski, & Jedrzewska, 2012;Petrunenko et al., 2016), in some cases, more so compared to gaur and muntjac (Sunquist et al., 1999;Biswas & Sankar, 2002). Hayward et al. (2012) suggest tigers optimally select prey with a similar body mass, corresponding roughly to large cervids (e.g. ...
... Wild boar and deer species, such as sambar, are suggested to be important prey species elsewhere in the tiger's range (Sunquist, Karanth, & Sunquist, 1999;Biswas & Sankar, 2002;Hayward, Jedrzejewski, & Jedrzewska, 2012;Petrunenko et al., 2016), in some cases, more so compared to gaur and muntjac (Sunquist et al., 1999;Biswas & Sankar, 2002). Hayward et al. (2012) suggest tigers optimally select prey with a similar body mass, corresponding roughly to large cervids (e.g. sambar) and wild boar which we considered 'large' and 'medium' prey species, respectively. ...
Thailand is one of the last strongholds for tigers Panthera tigris in mainland Southeast Asia. Evidence suggests heterogeneity in tiger presence in a globally important landscape in Eastern Thailand is potentially influenced by a complex interaction of prey, human presence and environmental conditions. Understanding these dynamics is of considerable importance for the conservation of tigers both in this landscape and elsewhere in their range. In this study, we examine which factors , among prey, human presence and environmental characteristics, best explain tiger presence in the Dong Phayayen-Khao Yai Forest Complex (DPKY). We collated survey data from 56,214 camera trap nights and evaluated the relationship between tiger presence and a suite of five prey, 11 human presence and eight environmental variables. We then used variance partitioning to discern the degree of variance in tiger presence explained by these factors. We documented strong, positive associations with wild boar Sus scrofa presence and prey richness, and strong, negative associations with human settlement density, public roads and presence of poachers. Environmental characteristics explained a greater relative proportion of variance (19.6%) in tiger presence than prey covariates alone (3.1%), particularly confounded with human presence (31.1%). This suggests that environmental variables , especially when accompanied by anthropogenic factors, could be used to model potential tiger occurrence where other data may be lacking. Our approach may be helpful in providing guidance for prioritizing habitat, evaluating the effect of human presence and identifying key prey to provide a foundation for tiger protection and recovery.
... Our objectives were to (1) document the minimum number of individual tigers in the area, (2) determine the temporal and spatial overlap of tigers with six potential prey species, and (3) create a composite score from the indices of temporal and spatial overlap as a novel method to predict predator-prey encounter rates and determine potential prey preference. In our analyses we included all potential tiger prey species present in the study area, based on a review of tiger dietary studies (Hayward et al., 2012). In line with known prey preferences of tigers across their range, we expected sambar deer Rusa unicolor and wild boar Sus scrofa to have the highest composite spatio-temporal score. ...
... We used kernel density estimation to determine activity patterns and quantify overlap among species (Ridout & Linkie, 2009). We reviewed potential prey species for tigers (Hayward et al., 2012) and analysed those in our study area with >75 detection events, which included the greater and lesser mouse deer (n=340), Malay tapir Tapirus indicus (n=85), pig-tailed macaque Macaca nemestrina (n=433), red muntjac Muntiacus muntjac (n=711), sambar deer (n=102), and wild boar (n=302). We first converted the time of each event into a radians measurement for analysis. ...
... Secondly, we considered a composite score that also included prey mass, with a higher mass adjustment value (spatial and temporal composite score´1.1) for prey within the preferred size range of tigers (60-250 kg; Hayward et al., 2012) and a lower value (spatial and temporal composite score´0.9) for potential prey outside this range. ...
The encounter rates carnivores have with prey are dependent on spatial and temporal overlap, and are often highest with their preferred prey. Sumatran tigers Panthera tigris sumatrae are critically endangered and dependent on prey populations, but little is known about their prey preferences. We collected 7 years of camera trapping data from Bukit Barisan Selatan National Park to investigate spatial and temporal overlap of tigers with potential prey species. We also developed a novel method to predict predator-prey encounter rates and potential prey preferences from camera trapping data. We documented a minimum of 10 individual tigers with overall relative abundance of 0.24 detections/100 trap nights for the population. Tigers exhibited a diurnal activity pattern and had highest temporal overlap with wild boar Sus scrofa (Δ1 = 0.80) and pigtail macaques Macaca nemestrina (Δ1 = 0.76), but highest spatial overlap with wild boar (AUC = 0.71) and sambar deer Rusa unicolor (AUC = 0.66). We created a spatial and temporal composite score and three additional composite scores for adjusted spatial weight and preferred prey mass, and each indicated tigers had the greatest overlap with wild boars followed by sambar deer, their known preferred prey in other areas. Spatial and temporal overlaps are often considered as separate indices, but a composite score may allow better predictions of encounter rates and potential prey preferences. Our findings suggest that prey management efforts in this area should focus on wild boar and sambar deer to ensure a robust prey base for this critically endangered tiger population.
... Availability of a sufficient prey base of large ungulates is the Tiger's primary habitat requirement: "wild pigs and deer of various species are the two prey types that make up the bulk of the Tiger's diet, and in general Tigers require a good population of these species to survive and reproduce" (Sunquist and Sunquist 2002). Tigers need to kill 50-60 large prey animals per year (Karanth et al. 2004, Hayward et al. 2012, Miller et al. 2013). However, when large prey populations are depleted they opportunistically predate on sub-optimal prey such as birds, fish, rodents, insects, amphibians, reptiles, and other mammals such as primates and porcupines. ...
... Tigers can also take ungulate prey much larger than themselves, including large bovids (Water Buffalo, Gaur, Banteng), and rarely even Asian Elephants and rhinos. However, like many large carnivores, preferred prey essential for successful reproduction are species that are approximately the same weight as Tigers themselves (Hayward et al. 2012). ...
Tiger Panthera tigris has most recently been assessed for The IUCN Red List of Threatened Species in 2021. Panthera tigris is listed as Endangered under criteria A2abcd.
... Therefore, wild pig and barking deer predation by both carnivores could be attributed to the abundance of the species or low abundance of medium-sized prey (chital) in the study area. Furthermore, our study conducted during the dry season, when wild pig mostly found solitary and more vulnerable to both ambush predators [56,57]. In Southwest Primorskii Krai, Russia, the wild pig was one of the important prey for both predators, where Amur tigers preyed on adult individuals and Amur leopards primarily preyed on smaller, sub-adult individuals [58]. ...
... Several studies reported the primate was one of the prime prey species for leopards in its range due to arboreal ability may allow them to take advantage of catching prey in the upper strata [55,60,61]. Following the previous study [57], tigers showed the preference for sambars and wild pigs in our study area. The high predation on these two species by tigers is likely to explain by their availability and body mass [23]. ...
There is a little understanding of how apex carnivores partition their diet to coexist. We studied food habit and trophic niche overlap of two apex carnivores, tiger and leopard in the tropical forest of Similipal tiger reserve, eastern India. We used line transect and scat analysis method, to estimate the prey availability and determine the diet and prey selection of two apex carnivores. Tigers consumed mostly large and medium-sized prey, whereas leopards mostly consumed medium and small-sized prey. Both carnivores were not random in their consumption of prey, sambar and wild pigs were selectively consumed by tigers, whereas leopards selectively consumed wild pigs, barking deer and mouse deer. Dietary overlap between two carnivore species was moderate (Pianka's niche overlap index: 0.55), and trophic niche breadth (Levin's standardized niche breadth) of the leopard (0.52) was greater than that of the tiger (0.37). Overall, tigers exhibited specialized feeding habits, whereas leopards showed generalist feeding habits. Our study highlights the opportunistic nature of leopards and probably a reason for the species successfully coexist with tigers.
... It is suggested that the density and spatial distribution of tigers are primarily determined by ungulate prey abundance (Karanth et al. 2004, Wegge and Storaas 2009, Harihar and Pandav 2012 and absence of human disturbances (Linkie et al. 2006 (Karanth et al. 2004, Wangchuk 2004, Harihar and Pandav 2012, Hayward et al. 2012, Tempa 2017. As such, these prey species will constitute an important structural composition of the tiger habitats & tiger movement corridors. ...
... Earlier studies reported sambar as one of the preferred prey species among the ungulate guilds (Johnsingh 1992, Karanth and Sunquist 1995, Bagchi et al. 2003 and similar findings were also reported from Bhutan (Wang 2008, Tempa 2017. However, in areas where large prey like sambar and gaur are absent, wild pigs and barking deer become important for tigers (Biswas and Sankar 2002, Hayward et al. 2012) so probability of using site A of the BC8 for movement of tiger between protected areas cannot be ruled out. Rather, this gives positive outlook as tigers in Bhutan prey upon wild pigs as evident from scats of tigers (Wang and Macdonald 2009). ...
The ability of animals to move across complex landscapes is critical for maintaining viable populations in the landscape. For an apex predator like tiger (Panthera tigris), which is territorial and requiring large home range, habitat connectivity through biological corridors is vital. In Bhutan, the landscape conservation approach for tiger was initiated since 1999 when biological corridors were first established. Yet, the status of connectivity are not widely known. This study assessed the structural connectivity of biological corridor number 8 (BC8) for the tiger movements between national parks considering the ecological and anthropogenic variables influencing tiger and its prey. Camera trap datasets were used to assess the single-season occupancy of three principal prey species of tiger, sambar (Rusa unicolor), barking deer (Muntiacus muntjak), and wild boar (Sus scrofa), and to determine the habitat use probability of tiger. A questionnaire survey was administered to assess the current trends of human-tiger conflict, and people's perception towards conservation. Barking deer had the highest occupancy estimate (ψ ± SE) 0.52 ± 0.09, followed by sambar (ψ ± SE) 0.49 ± 0.03 and wild boar (ψ ± SE) 0.45 ± 0.07. The influence of ecological and anthropogenic variables like elevation, aspect, slope, distance to river, and distance to settlement varied among the three prey species. Predicted occupancy map showed probable niche partitioning between species, thereby enabling better distribution of principal prey and thus the connectivity. Tiger habitat use probability was influenced positively by elevation and negatively by aspect and slope. Livestock predation was the prevailing human-tiger conflict in the area with tiger accounting 76.49% of the total kill in the past two years. Nomads were more vulnerable than agro-pastoralists, and livestock predation was higher during the winter. Over 80% of the respondents were not aware of biological corridors and over 30% perceived negative attitude towards tiger conservation and corridor management. The structural connectivity maintained by ecological variables and prey species would enable tiger movements across the BC8, but human-tiger conflict needs to be mitigated, and people should be educated for conservation. Management of the BC8 by developing the management plan remains the most pressing need.
... These weights are key to estimating the biomass of these species in the diet of tigers. Because adult male gaur are 1.3 times larger than adult male banteng, and males of both species are >3.5 times larger than Hayward et al. (2012) optimum 1:1 predator to prey ratio, we hypothesize that these species, and especially the large size classes, are approaching the maximum size limit of tiger prey. Support for this hypothesis would be (a) male tigers kill a smaller percentage of larger adult male gaur than smaller adult banteng, (b) female tigers, which average two thirds the weight of adult male tigers, kill a smaller percentage of adult males of both species compared with male tigers, and (c) female tigers kill fewer males of both species compared with males. ...
... Support for this hypothesis would be (a) male tigers kill a smaller percentage of larger adult male gaur than smaller adult banteng, (b) female tigers, which average two thirds the weight of adult male tigers, kill a smaller percentage of adult males of both species compared with male tigers, and (c) female tigers kill fewer males of both species compared with males. It is important to note that our research does not imply prey preference (Hayward et al., 2012); we simply report size, sex, and age class of gaur and banteng killed by tigers in our study area. ...
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.
... The body size, availability and vulnerability of prey are the primary factors determining prey selection. A recent metaanalysis (Hayward et al. 2012) suggests that species weighing between 60 and 250 kg are preferred by tigers, and in our study the two large prey, wild boar (103 kg) and sika deer (95 kg) were the preferred prey. The wild boar is the largest ungulate preferred by Amur tigers (Fig. 3), supporting the findings of Hayward et al. (2012); they report that the wild boar is one of the species most preferred by tigers based on selectivity index scores from 3187 kills or scats from 32 prey species. ...
... A recent metaanalysis (Hayward et al. 2012) suggests that species weighing between 60 and 250 kg are preferred by tigers, and in our study the two large prey, wild boar (103 kg) and sika deer (95 kg) were the preferred prey. The wild boar is the largest ungulate preferred by Amur tigers (Fig. 3), supporting the findings of Hayward et al. (2012); they report that the wild boar is one of the species most preferred by tigers based on selectivity index scores from 3187 kills or scats from 32 prey species. Our results are also congruent with those of earlier studies from reserves in Russia (Kerley and Borisenko 2007, Kerley et al. 2015, Sugimoto et al. 2016). ...
... Secara global, rusa sambar adalah spesies mangsa yang dominan dari segi biojisim dan diutamakan dalam diet harimau sepanjang masa (Biswas & Sankar 2002;Hayward, Jedrzejewski & Jedrzewska 2012). Harimau adalah felid terbesar dan memakan ungulat besar hampir keseluruhannya. ...
Populasi rusa sambar (Rusa unicolor) mengalami penurunan dan dikategorikan sebagai rentan oleh Kesatuan Antarabangsa untuk Pemuliharaan Alam dan Sumber Asli pada tahun 2014. Objektif kajian ini untuk mengenal pasti faktor sekitaran yang mempengaruhi kesesuaian habitat rusa sambar, yang terdiri daripada kecerunan, ketinggian, guna tanah, jarak daripada bandar, jarak daripada sungai dan litologi dianalisis menggunakan sistem maklumat geografi (GIS). Data kehadiran rusa sambar dari tahun 1992-2008 diintegrasikan dengan faktor sekitaran menggunakan model entropi maksimum untuk mengenal pasti habitat yang sesuai bagi rusa sambar di kompleks hutan utama Taman Negara. Hasil kajian ini menunjukkan jarak dari sungai menyumbang peratusan paling tinggi dalam model kesesuaian habitat iaitu 47.7%. Manakala jarak dari bandar, jenis guna tanah, ketinggian, kecerunan dan litologi menyumbang sebanyak 21.8%, 16.1%, 7.2%, 6.7% dan 0.4%. Habitat yang sesuai bagi rusa sambar meliputi 37% daripada keseluruhan kawasan kajian (2,218,389 hektar). Kajian penentuan potensi habitat rusa sambar yang sesuai di kompleks hutan utama Taman Negara penting untuk pengurusan hidupan liar seperti aktiviti pelepasan semula.
... gregarious muntjac deer (20+ individuals per sampling occasion), we followedBrodie et al. (2018) by analyzing the number of muntjac groups rather than the number of individuals in the count history and limited the daily observations to zero or one group. A systematic range-wide dietary study suggests tigers preferentially prey upon the largest available prey species with a weight range from 60 to 250 kg, such as sambar deer(Hayward et al., 2012), while single-landscape studies highlight muntjac deer as an important prey species for clouded leopards(Can et al., 2020; ...
Predator–prey dynamics are a fundamental part of ecology, but directly studying interactions has proven difficult. The proliferation of camera trapping has enabled the collection of large datasets on wildlife, but researchers face hurdles inferring interactions from observational data. Recent advances in hierarchical co‐abundance models infer species interactions while accounting for two species' detection probabilities, shared responses to environmental covariates, and propagate uncertainty throughout the entire modeling process. However, current approaches remain unsuitable for interacting species whose natural densities differ by an order of magnitude and have contrasting detection probabilities, such as predator–prey interactions, which introduce zero inflation and overdispersion in count histories. Here, we developed a Bayesian hierarchical N‐mixture co‐abundance model that is suitable for inferring predator–prey interactions. We accounted for excessive zeros in count histories using an informed zero‐inflated Poisson distribution in the abundance formula and accounted for overdispersion in count histories by including a random effect per sampling unit and sampling occasion in the detection probability formula. We demonstrate that models with these modifications outperform alternative approaches, improve model goodness‐of‐fit, and overcome parameter convergence failures. We highlight its utility using 20 camera trapping datasets from 10 tropical forest landscapes in Southeast Asia and estimate four predator–prey relationships between tigers, clouded leopards, and muntjac and sambar deer. Tigers had a negative effect on muntjac abundance, providing support for top‐down regulation, while clouded leopards had a positive effect on muntjac and sambar deer, likely driven by shared responses to unmodelled covariates like hunting. This Bayesian co‐abundance modeling approach to quantify predator–prey relationships is widely applicable across species, ecosystems, and sampling approaches and may be useful in forecasting cascading impacts following widespread predator declines. Taken together, this approach facilitates a nuanced and mechanistic understanding of food‐web ecology. Predator–prey dynamics are a fundamental part of ecology, but studying interactions from observational data (e.g., camera trapping) has proven difficult. Here, we developed a Bayesian hierarchical N‐mixture co‐abundance model that is suitable for inferring predator–prey interactions by accounting for excessive zeros and overdispersion in count histories. Our co‐abundance model detected clear positive and negative predator–prey relationships from a large Southeast Asian camera trapping dataset, and we highlight how it overcomes barriers, is widely applicable across species, ecosystems, and sampling approaches, and may be useful in forecasting cascading impacts following widespread predator declines.
... Other authors have speculated that the capacity for deer to cope physiologically with being pursued by hounds is shaped by their evolutionary or individual histories (Bateson and Bradshaw 1997). In this context, sambar deer have evolved with a suite of large mammalian predators, such as, for example, tigers (Panthera tigris) in southern Asia (Hayward et al. 2012) and are likely to be harrassed or attacked occasionally by dingoes/wild dogs in contemporary southeastern Australia (Forsyth et al. 2019). Although the stress experienced prior to death is likely to be significant for sambar deer hunted with hounds, the impact of hunting on chronic stress of surviving deer is unknown (Sauerwein et al. 2004). ...
... 9: 220697 the vastly different environmental conditions of the Russian Far East, compared with the rest of the continental tiger range, which have promoted greater frequency and magnitude of forces acting on the skull and mandible in killing prey and chewing. The prey composition and preferences of the Amur population could have led to skull differences, yet large deer species, similar in body size to the red deer found in the Russian Far East, are preferentially preyed upon in southern continental tiger populations, and wild pigs, Sus scrofa, are a ubiquitous staple of tiger diets [19,56,57]. We propose two possible scenarios for increased jaw musculature in wild Amur tigers compared with other wild continental tigers that have resulted in the skull morphological changes reported here: Amur tigers may well have to chew through frozen carcasses in winter when they stay beside a kill until it is consumed, requiring increased masticatory forces when compared with other continental tigers. ...
Tiger subspecific taxonomy is controversial because of morphological and genetic variation found between now fragmented populations, yet the extent to which phenotypic plasticity or genetic variation affects phenotypes of putative tiger subspecies has not been explicitly addressed. In order to assess the role of phenotypic plasticity in determining skull variation, we compared skull morphology among continental tigers from zoos and the wild. In turn, we examine continental tiger skulls from across their wild range, to evaluate how the different environmental conditions experienced by individuals in the wild can influence morphological variation. Fifty-seven measurements from 172 specimens were used to analyse size and shape differences among wild and captive continental tiger skulls. Captive specimens have broader skulls, and shorter rostral depths and mandible heights than wild specimens. In addition, sagittal crest size is larger in wild Amur tigers compared with those from captivity, and it is larger in wild Amur tigers compared with other wild continental tigers. The degree of phenotypic plasticity shown by the sagittal crest, skull width and rostral height suggests that the distinctive shape of Amur tiger skulls compared with that of other continental tigers is mostly a phenotypically plastic response to differences in their environments.
... Like other large, solitary predators, tigers (Panthera tigris) and leopards (P. pardus), snow leopards may need to be more selective about the habitat zone they forage in and the weight class of prey they target in order to maximize hunting success and reduce injury risk [47,49]. Indeed, it is likely that snow leopards, like lions (P. ...
... • E and 42.614041 • N-43.553505 • N) (Figure 1), all of which followed the same pattern: the material for building the nest was obtained by biting off the bushes around the nest (Figure 1, Video S1). This pattern is perplexing; when wild boars use the nest, they also choose to aggressively expose themselves, increasing the risk of predation by the Amur tiger, which prefers to be as close to the prey as possible through ambush behavior to enhance prey 'catchability' [35]. The Amur tiger-wild boar relationship is a typical predation and anti-predation interaction dependent on vision. ...
Exploring the processes of interspecific relationships is crucial to understanding the mechanisms of biodiversity maintenance. Visually detecting interspecies relationships of large mammals is limited by the reconstruction accuracy of the environmental structure and the timely detection of animal behavior. Hence, we used backpack laser scanning (BLS) to reconstruct the high-resolution three-dimensional environmental structure to simulate the process of a predator approaching its prey, indicating that predator tigers would reduce their visibility by changing their behavior. Wild boars will nibble off about 5m of branches around the nest in order to create better visibility around the nest, adopting an anti-predation strategy to detect possible predators in advance. Our study not only points out how predator–prey relationships are affected by visibility as the environment mediates it, but also provides an operable framework for exploring interspecific relationships from a more complex dimension. Finally, this study provides a new perspective for exploring the mechanisms of biodiversity maintenance.
... B 289: 20212681 and weaklings or scavenge on tiger kill carcasses [64]. Furthermore, it is possible that in a prey-rich environment, tigers may prefer large-bodied prey sparing smaller prey species for leopards thus segregating along the diet niche [65,66]. Our findings highlight that abundant large prey is critical for supporting large carnivore communities [67]. ...
In the face of a growing human footprint, understanding interactions among threatened large carnivores is fundamental to effectively mitigating anthro- pogenic threats and managing species. Using data from a large-scale camera trap survey, we investigated the effects of environmental and anthropogenic variables on the interspecific interaction of a carnivore guild comprising of tiger, leopard and dhole in Bhutan. We demonstrate the complex effects of human settlement density on large carnivore interactions. Specifically, we demonstrate that leopard–dhole co-occupancy probability was higher in areas with higher human settlement density. The opposite was true for tiger–leopard co-occupancy probability, but it was positively affected by large prey (gaur) abundance. These findings suggest that multi-carnivore communities across land-use gradients are spatially structured and mediated also by human presence and/or the availability of natural prey. Our findings show that space-use patterns are driven by a combination of the behavioural mechanism of each species and its interactions with competing species. The duality of the effect of settlement density on species interactions suggests that the benefits of exploiting anthropogenic environments are a trade-off between ecological opportunity (food subsidies or easy prey) and the risk of escalating conflict with humans.
... The tiger exhibited only random associations with small prey like barking deer, while the leopard had positive associations. Large-bodied wild ungulates such as the sambar, wild boar, gaur, and chital are a chief constituent of the tiger diet (Mondal et al. 2011;Hayward et al. 2012;Basak et al. 2020). The proportion of the domestic prey is almost zero in the tiger diet, and the buffalo hairs observed in the scat can be feral or free-ranging. ...
Tiger conservation is a global initiative, and data on distribution, prey dynamics, competition, and range extension are critical factors in sustaining its fragile populations. We concentrated on evaluating the data pertaining to these variables in order to designate the high-altitude Nilgiri forest division as a tiger conservation unit. We gathered secondary data on animal density, mortality, and conflicts. We also conducted a people perception survey, a systematic grid-based prey species survey, and a tiger and leopard scat survey to assess the prey-predator relationship and understand human attitudes toward carnivore conservation. According to the findings, the Nilgiri forest division has a healthy prey base with positive or random prey-predator associations and a significant correlation between mammalian assemblages. Because the niche overlap between the tiger and the leopard is high, the latter broadens its niche and relies on wild prey in the shola fringes and tea estates. The tiger avoids human-dominated areas and prefers to stay in the shola, rarely venturing into tea estates. In contrast to previous considerations, we believe the Nilgiri forest division is an ideal tiger habitat. We specify that instead of being considered a connective corridor, the Nilgiri forest division may be merged with Mukkurthi National Park to form a high-altitude tiger reserve.
... Numbers close to zero indicate resource use in proportion to availability. This index has been used on many other studies to establish prey preference by predators (Hayward et al., 2006(Hayward et al., , 2012Nowak et al., 2011;Lyngdoh et al., 2014). ...
ABSTRACT—Retaliatory killing of large carnivores due to livestock predation is one of the major threats for the conservation of many declining populations of predators. According to empirical observations, there is a higher incidence of livestock predation when native prey abundance is low. In this study, we applied a treatment consisting of augmentation of prey abundance by translocation of peccaries (Pecari tajacu) and placement of four feed stations for white-tailed deer (Odocoileus virginianus) on a cattle ranch in Sonora, Mexico, with verified calf predation by puma (Puma concolor) and jaguar (Panthera onca). We quantified and compared consumed prey over two periods—phase I (8 months before the augmentation of prey) and phase II (8 months after the augmentation of prey)—through investigation of kill sites from Global Positioning System–collared jaguar and puma, prey identification from analyzed scat using molecular DNA techniques, and opportunistic discoveries of recently killed animal remains by either predator. We calculated the relative abundance of species (17 mammals [one species with two distinct age classes] and 1 bird species) through camera traps and for the most relevant prey species for this study (deer, calf, and peccary), we also estimated prey use by the predator, based on their availability during each period (prey preference). In the prey composition analyses of scat, we observed a significant reduction in the consumption of bovids and a significant increase in the consumption of peccaries during phase II. In the analyses of prey use, during phase I, predators consumed peccaries and calves at a higher proportion in relation to their availability. During phase II, consumption of calves declined from being preferred, to being consumed at the same proportion as their availability. Application of these results can contribute to the decrease of livestock predation and therefore conservation of pumas and jaguars.
... This study was part of a larger effort to estimate habitat use of wild carnivores and herbivores within the agricultural corridor. Wild boar are an important tiger prey species (Hayward et al., 2012) and are widely regarded as an agricultural pest across their distributional range (Lewis et al., 2017). We explored how conflicts with wild boar are reported from sites occupied and unoccupied by the species. ...
Managing human-wildlife conflicts (HWCs) is an important conservation objective for the many terrestrial landscapes dominated by humans. Forecasting where future conflicts are likely to occur and assessing risks to lives and livelihoods posed by wildlife are central to informing HWC management strategies. Existing assessments of the spatial occurrence patterns of HWC are based on either understanding spatial patterns of past conflicts or patterns of species distribution. In the former case, the absence of conflicts at a site cannot be attributed to the absence of the species. In the latter case, the presence of a species may not be an accurate measure of the probability of conflict occurrence. We present a Bayesian hierarchical modeling framework that integrates conflict reporting data and species distribution data, thus allowing the estimation of the probability that conflicts with a species are reported from a site, conditional on the species being present. In doing so, our model corrects for both false-positive and false-negative conflict reporting errors. We provide study design recommendations using simulations that explore the performance of the model under a range of conflict reporting probabilities. We applied the model to data on wild boar (Sus scrofa) space use and conflicts collected from the Central Terai Landscape (CTL), an important tiger conservation landscape in India. We found that tolerance for wildlife was a predictor of the probability with which farmers report conflict with wild boars from sites not used by the species. We also discuss useful extensions of the model when conflict data are verified for potential false-positive errors and when landscapes are monitored over multiple seasons.
... The preferred prey size class is around the same weight as the tiger itself. Per research by Hayward et al. 2012, significantlypreferred prey are sambar and wild boar, with significantly-avoided species being chital, chinkara, nilgai, macaque, langur, and peafowl (Hayward et al., 2012). Tigers generally require >5 kg meat daily to maintain body condition (Sunquist, 1981). ...
As the human population increases and climate change exacerbates resource scarcity, India's wildlife faces the increasing threat of fragmentation and habitat loss in a human-centric landscape. The wild exists in small semi-isolated pockets, connected sparsely, if at all, by wildlife corridors. Such wildlife corridors may be the last hope for charismatic megafauna such as the Bengal tiger (Panthera tigris tigris). This project assesses the impacts of encroachment and land use change on the use of two wildlife dispersal (corridors) connecting Ranthambhore National Park (RNP) to other intact forest patches in the larger Ranthambhore Tiger Reserve (RTR) by the tiger and other large carnivores. The research objectives are: (1) to quantify land use and land cover change between 2011 and 2016 in RTR; (2) to assess the effects of this change on the two major wildlife dispersal corridors; and (3) to better understand how this affects land cover preference and use by tigers. Field data collection of wildlife presence and human encroachment in the two corridors was carried out during May-June 2016, and geospatial analysis was used to generate maps for WWF-India, as well as to study the changes in the landscape over the five-year period. Major threats to corridors were found to be the expansion of agricultural land, sand mining, overgrazing of forest vegetation by livestock, and urban expansion. Agricultural expansion has caused a decline in forest and ravine land cover, which are preferred by dispersing carnivores. Most threats were concentrated around the northwest corridor, making it a conservation priority. The southeast corridor is also heavily used by dispersing tigers as it is mostly characterized by ravines, a preferred habitat type; however, flattening of ravines for agricultural expansion and mining brings carnivores into proximity with humans, increasing the potential for conflict in the region. Based on these results, we recommend increased protection of the buffer zone of the northwest corridor and suggest that remaining efforts be put towards conservation education and stricter regulation of land use practices in areas surrounding corridors and intact habitat patches.
... Population densities of large predators are closely tied to the biomass or density of prey (Hayward et al., 2007;Karanth et al., 2004;Miquelle et al., 2010). As tigers prefer medium and large sized prey (Hayward et al., 2012;Miquelle et al., 2010) and their successful reproduction is likely dependent on adequate densities of these preferred species (Miller et al., 2014;Miquelle et al., 2018), our estimates of tiger carrying capacity based on total number of prey is considered overly simplistic and an overestimate. In contrast, the estimates of tiger carrying capacity adjusted for relative availability among different prey species are therefore, likely more realistic as a basis for estimating present carrying capacity. ...
With most large carnivore populations and habitats suffering massive declines across the world, efforts to restore their populations in the wild are of high conservation priority. China attaches great importance to protecting Amur tigers and has made considerable progress towards this goal, but most efforts have been focused on the single Laoyeling landscape. Here we assess all four major forested landscapes (including Laoyeling, Zhangguangcailing, Wandashan and Lesser Khinghan Mountains) that may be suitable for tigers in northeast China. Between 2013 and 2018, in total 54 wild Amur tigers were recorded through camera trap surveys and 30 individuals were identified through genetic analyses. The finite rate of increase of the population in the Laoyeling landscape, the only landscape with a breeding population, fluctuated greatly, but averaged 1.51. Total habitat used by Amur tigers during this period amounted to 47,813 km². We estimated that based on density and home range sizes in nearby Russia, these landscapes could support 311 tigers, including 119 resident breeding females. However, prey density, especially of preferred prey, was extremely low, and likely a primary constraint to population recovery in landscapes other than Laoyeling, especially the Wandashan and Lesser Khingan Mountains landscapes. To achieve the goal of a large meta-population of Amur tigers across northeast Asia, in addition to prey restoration, we recommend extensive landuse planning, reducing anthropogenic impacts, improving ecology connectivity, and extensive international cooperation, as well as a feasibility assessment for introducing orphaned cubs as a means of speeding recovery in landscapes without breeding females.
... Hunting for meat is an important evolutionary mechanism for driving larger brain sizes and innovation in hominids (Aiello & Wheeler, 1995). We conclude that chimpanzees are not apex predators of vertebrates in the way that modern humans, lions (Panthera leo), and tigers (Panthera tigris) are apex predators (Hayward et al., 2012;Hayward & Kerley, 2005). ...
The common chimpanzee Pan troglodytes is the closest extant relative of modern humans and is often used as a model organism to help understand prehistoric human behavior and ecology. Originally presumed herbivorous, chimpanzees have been observed hunting 24 species of birds, ungulates, rodents, and other primates, using an array of techniques from tools to group cooperation. Using the literature on chimpanzee hunting behavior and diet from 13 studies, we aimed to determine the prey preferences of chimpanzees. We extracted data on prey‐specific variables such as targeted species, their body weight, and their abundance within the prey community, and hunter‐specific variables such as hunting method, and chimpanzee group size and sex ratio. We used these data in a generalized linear model to determine what factors drive chimpanzee prey preference. We calculated a Jacobs’ index value for each prey species killed at two sites in Uganda and two sites in Tanzania. Chimpanzees prefer prey with a body weight of 7.6 ± 0.4 kg or less, which corresponds to animals such as juvenile bushbuck (Tragelaphus scriptus) and adult ashy red colobus monkeys (Piliocolobus tephrosceles). Sex ratio in chimpanzee groups is a main driver in developing these preferences, where chimpanzees increasingly prefer prey when in proportionally male‐dominated groups. Prey preference information from chimpanzee research can assist conservation management programs by identifying key prey species to manage, as well as contribute to a better understanding of the evolution of human hunting behavior.
... This study showed that the animal most affected by livestock was sika deer, the dominant prey in the diet of tigers and leopards (Kerley et al. 2015;Sugimoto et al. 2016). Given that availability of ungulate prey is crucial to the persistence of any tiger and leopard population (especially large-bodied deer for tigers) (Karanth et al. 2004;Hayward et al. 2012;Li et al. 2019), the results of this study have conservation implications in terms of assessing the cascading effects of cattle grazing through a multispecies perspective (Filazzola et al. 2020). Tigers and leopards are now showing a trend towards expanding their range into China (Wang et al. 2015;Ning et al. 2019) but with distribution still spotty and overall numbers low (Vitkalova et al. 2018;Wang et al. 2018). ...
ContextLivestock grazing is one of the most widespread types of anthropogenic land use, even occurs in many protected areas and has become a threat to wildlife worldwide. Understanding livestock-wildlife interactions is crucial for rare large carnivores conservation. In China, free-ranging cattle within forests degrade the habitat of the tigers (Panthera tigris) and leopards (Panthera pardus), but quantitative assessments of how livestock affect the spatial and temporal use by the major ungulate prey of the two endangered felids are very limited.Objectives
This study aimed to examine the interactions of several sympatric wildlife species with livestock at a fine spatiotemporal scale in a human-dominated forest landscape.Methods
Based on a large-scale camera-trapping data across the China-Russia border, we used N-mixture models, two-species occupancy models and activity pattern overlap to understand the effects of cattle grazing on three ungulate species (sika deer Cervus nippon, wild boars Sus scrofa and roe deer Capreolus pygargus).ResultsSpatially, with cattle activity increasing, wild boar and roe deer had different degrees of decline in the intensity of habitat use. Sika deer were displaced as more cattle encroached on forest habitat. Temporally, in the presence of cattle, wild boar and sika deer decreased their activities in the day. In addition, three wild ungulates trend to exhibit lower spatiotemporal overlap with cattle at shared camera sites.Conclusions
Our study shows that wildlife species may reduce the probability of habitat use by spatial avoidance and changing the daily activity patterns. We underscore that fine-scale (i.e. camera-site level) spatiotemporal avoidance is likely a key component of co-occurrence between livestock and the sympatry of competing ungulates inhabiting forest ecosystems. Given prey were depressed, efforts to minimize the livestock disturbance on these species need to be considered to ensure their sustained recoveries.
... Tigers may use wide and established trails for movement (Kawanishi, 2002), but leopards may avoid them by using smaller trails or different activity patterns. Our study showed considerably higher activity overlaps (>0.50) of tigers with medium-to large-sized prey (Hayward et al., 2012) and leopards with small-medium-to large-sized prey (Hayward et al., 2006). ...
Species interactions play a vital role in structuring mammalian communities by stimulating behavioral responses in varied niche dimensions that affect sympatric associations and predator–prey relationships. We determined temporal overlap and effects of the moon cycle on dominant and sub-dominant mammalian assemblages in Manas National Park, India. A total of 36 species were captured, with 24,865 independent records over 11,294 trap nights. We collected 1,130 photographs of five large- and medium-sized carnivores and 1,541 photographs of 12 small carnivores. Fifty-one percent of records were detected during diurnal period, followed by 38% in nocturnal phase, and 11% during twilight. Small carnivores such as Prionailurus bengalensis and Viverridae spp. were strictly nocturnal, whereas Martes flavigula and Herpestidae spp. were diurnal. Medium-sized carnivores were either nocturnal (Neofelis nebulosa) or diurnal (Cuon alpinus), whereas large-sized carnivores (Panthera tigris, Panthera pardus, and Ursus thibetanus) were cathemeral. A high degree of temporal overlap (>0.75) was found between most sympatric carnivores with distinct activity peaks, while a low overlap (<0.50) was observed between different body-sized carnivores. Viverrids’ activity was negatively correlated (r = −0.44, p < 0.01) with lunar cycles, perhaps to increase foraging efficiency or as an anti-predator strategy. Large prey (μ = 133.23°) and small prey (μ = 131.35°) activity were high during brighter nights due to better visual detection in detecting or avoiding predators. Dominant species activity was least affected by the lunar cycle among forest-dependent mammals, whereas subdominant species activity was either lunarphobic or lunarphilic. The study demonstrates the use of passive camera traps in understanding the behavioral rhythms of tropical mammals.
... With eight individuals positively recorded, the present study further confirms than JSWNP is an important tiger conservation landscape in Bhutan with the potential to serve tiger source population (UWICE, 2015) for further dispersal to northern protected areas of the country. The presence of many ungulates co-occurring with tiger further gives us a posit that easy availability of prey species will enable favourable conditions for tiger to persist (Karanth et al., 2004;Hayward et al., 2012). Ungulates like sambar, wild boar, barking deer, chital, and gaur, are considered preferred prey of the tiger in the Himalayan landscape depending on locality and habitat type (Karanth et al., 2004;Harihar & Pandav, 2012;Tempa, 2017)ecologists should be able to identify a small number of limiting resources for a species of interest, estimate densities of these resources at different locations across the landscape, and then use these estimates to predict the density of the focal species at these locations. ...
This report presents the findings from two consecutive monitoring surveys (2016-2017
(2017), and 2017-2018 (2018)). We report the number of tiger individuals captured in
JSWNP during the two surveys, the diversity of wild felids and its sympatry with tigers,
and occupancy and habitat use estimates of tiger and its prey species in the national
park. We used remotely triggered camera traps to photo capture tigers of JSWNP in the
potential tiger habitats. We established 39 camera stations during the first monitoring
and 41 stations during the second monitoring session, with each camera trap functioning
at least 60 days, over a total of 2617 and 2819 camera trap days respectively.
... While most solitary large felids prefer prey similar to their own size or smaller (e.g. Hayward et al. 2006Hayward et al. , 2012, snow leopards predominantly hunt ibex, blue sheep (Pseudois nayaur) and Himalayan Tahr (Hemitragus jemlahicus) (Lovari et al. 2013;Lyngdoh et al. 2014), which are considerably heavier (mean weight 55 kg; Lyngdoh et al. 2014) than snow leopards (mean weight 40 kg; Johansson et al. 2015). Because of the mountainous habitat occupied by the species and the large prey relative to predator size, opportunities for snow leopard mothers to release large prey for the cubs to successfully practice killing is rare (e.g. in contrast to cheetahs; Caro 1994, and tigers;Chundawat 2018). ...
Significant knowledge gaps persist on snow leopard demography and reproductive behavior. From a GPS-collared population in Mongolia, we estimated the timing of mating, parturition and independence. Based on three mother-cub pairs, we describe the separation phase of the cub from its mother as it gains independence. Snow leopards mated from January-March and gave birth from April-June. Cubs remained with their mother until their second winter (20-22 months of age) when cubs started showing movements away from their mother for days at a time. This initiation of independence appeared to coincide with their mother mating with the territorial male. Two female cubs remained in their mothers' territory for several months after initial separation, whereas the male cub quickly dispersed. By comparing the relationship between body size and age of independence across 11 solitary, medium-to-large felid species, it was clear that snow leopards have a delayed timing of separation compared to other species. We suggest this may be related to their mating behavior and the difficulty of the habitat and prey capture for juvenile snow leopards. Our results, while limited, provide empirical estimates for understanding snow leopard ecology and for parameterizing population models.
... Excluding placental (subfamilies Talpinae and Scalopinae) and marsupial moles (Notoryctes spp.) and monotremes, all terrestrial mammalian species have visible pinnae (Aitkin, 1997). The pinnae amplifies and transfers information (through sound reflection) to the middle and inner ear for interpretation by the brain (Hayward, Jędrzejewski, & Jêdrzejewska, 2012;Rosowski, 1996). Removal experiments, such as those conducted on the insectivorous brown long-eared bat (Plecotus auritus) (Muller, Lu, & Buck, 2008), that has large prominent pinnae, have provided information about the performance of information intake by the pinnae. ...
Multiple studies have described the anatomy and function of the external ear (pinna) of bats, and other placental mammals, however, studies of marsupial pinna are largely absent. In bats, the tragus appears to be especially important for locating and capturing insect prey. In this study, we aimed to investigate the pinnae of Australian marsupials, with a focus on the presence/absence of tragi and how they may relate to diet. We investigated 23 Australian marsupial species with varying diets. The pinnae measurements (scapha width, scapha length) and tragi (where present) were measured. The interaural distance and body length were also recorded for each individual. Results indicated that all nectarivorous, carnivorous, and insectivorous species had tragi with the exception of the insectivorous striped possum (Dactylopsila trivirgata), numbat (Myrmecobius fasciatus), and nectarivorous sugar glider (Petaurus breviceps). No herbivorous or omnivorous species had tragi. Based on the findings in this study, and those conducted on placental mammals, we suggest marsupials use tragi in a similar way to placentals to locate and target insectivorous prey. The Tasmanian devil (Sarcophilus harrisii) displayed the largest interaural distance that likely aids in better localization and origin of noise associated with prey detection. In contrast, the smallest interaural distance was exhibited by a macropod. Previous studies have suggested the hearing of macropods is especially adapted to detect warnings of predators made by conspecifics. While the data in this study demonstrate a diversity in pinnae among marsupials, including presence and absence of tragi, it suggests that there is a correlation between pinna structure and diet choice among marsupials. A future study should investigate a larger number of individuals and species and include marsupials from Papua New Guinea, and Central and South America as a comparison.
... Grassland and sparse vegetation are important components of the landscape for wild ungulates, small mammals, and birds, as well as for livestock, on which snow leopards are known to occasionally prey (Bagchi & Mishra, 2006;Chen et al., 2016;Lyngdoh et al., 2014). Therefore, the selection of a landscape with a total optimum amount of sparse vegetation in a given radius (as highlighted by PLAND_Gr), regardless of its extensiveness or continuity (revealed by GYR_AM_Gr) might be indicative of habitat choices intended to maximize hunting opportunities by foraging in preys' feeding grounds, and to balance tradeoffs in energy expenditures to locate and chase them (Hayward et al., 2011;Hayward, Jędrzejewski, & Jêdrzejewska, 2012;Lyngdoh et al., 2014). ...
Replicated multiple scale species distribution models (SDMs) have become increasingly important to identify the correct variables determining species distribution and their influences on ecological responses. This study explores multi‐scale habitat relationships of the snow leopard (Panthera uncia ) in two study areas on the Qinghai–Tibetan Plateau of western China. Our primary objectives were to evaluate the degree to which snow leopard habitat relationships, expressed by predictors, scales of response, and magnitude of effects, were consistent across study areas or locally landcape‐specific. We coupled univariate scale optimization and the maximum entropy algorithm to produce multivariate SDMs, inferring the relative suitability for the species by ensembling top performing models. We optimized the SDMs based on average omission rate across the top models and ensembles’ overlap with a simulated reference model. Comparison of SDMs in the two study areas highlighted landscape‐specific responses to limiting factors. These were dependent on the effects of the hydrological network, anthropogenic features, topographic complexity, and the heterogeneity of the landcover patch mosaic. Overall, even accounting for specific local differences, we found general landscape attributes associated with snow leopard ecological requirements, consisting of a positive association with uplands and ridges, aggregated low‐contrast landscapes, and large extents of grassy and herbaceous vegetation. As a means to evaluate the performance of two bias correction methods, we explored their effects on three datasets showing a range of bias intensities. The performance of corrections depends on the bias intensity; however, density kernels offered a reliable correction strategy under all circumstances. This study reveals the multi‐scale response of snow leopards to environmental attributes and confirms the role of meta‐replicated study designs for the identification of spatially varying limiting factors. Furthermore, this study makes important contributions to the ongoing discussion about the best approaches for sampling bias correction.
... Tigers inhabiting the mangrove forests of the Sundarbans prey upon a limited number of species of medium-sized ungulates compared to the wider range of medium-to large-sized ungulates that commonly occur in most other tiger landscapes (Karanth et al. 1995, Wang & Macdonald 2009, Miquelle et al. 2010, Hayward et al. 2012. Several large ungulate species, including wild water buffalo Bubalus arnee, ABSTRACT: The Sundarbans is the only mangrove habitat in the world to support tigers Panthera tigris, whose persistence there is believed to be dependent on a very limited number of prey species. ...
The Sundarbans is the only mangrove habitat in the world to support tigers Panthera tigris , whose persistence there is believed to be dependent on a very limited number of prey species. Conservation managers therefore need to understand how tigers utilise available prey species on a spatial scale in order to formulate a prey-based protection strategy for this global-priority tiger landscape. A total of 512 scat samples were collected during a survey of 1984 km ² of forest across 4 sample blocks in the 6017 km ² of the Bangladesh Sundarbans. Analysis of scat composition and prey remains reliably identified 5 major prey species, of which spotted deer Axis axis and wild pig Sus scrofa contributed a cumulative biomass of 89% to tiger diet. Tiger preference for prey species was highly skewed towards spotted deer and wild pig, but the relative contribution of these 2 species differed significantly across the 4 study areas, which spanned the Sundarbans, demonstrating important spatial patterns of tiger prey preference across the Sundarbans landscape. Given the comparatively limited number of prey species available to support the dwindling tiger population, different strategies are needed in different parts of the Sundarbans to support tiger populations and to protect spotted deer and wild pig populations from unabated poaching.
... Among large species of potential prey, wild pig was most widespread, followed by Indochinese serow. Wild pig is overall the most preferred prey of tiger (Hayward et al., 2012), and wild pig remained widely distributed in NEPL, which further supports our conclusion that tiger became extirpated because of the increase in snaring (Johnson et al., 2016), rather than for other reasons such as prey declines. ...
The Nam Et - Phou Louey National Protected Area (NEPL) is known for its diverse community of carnivores, and a decade ago was identified as an important source site for tiger conservation in Southeast Asia. However, there are reasons for concern that the status of this high priority diverse community has deteriorated, making the need for updated information urgent. This study assesses the current diversity of mammals and birds in NEPL, based on camera trap surveys from 2013 to 2017, facilitating an assessment of protected area management to date. We implemented a dynamic multispecies occupancy model fit in a Bayesian framework to reveal community and species occupancy and diversity. We detected 43 different mammal and bird species, but failed to detect leopard Panthera pardus and only detected two individual tigers Panthera tigris, both in 2013, suggesting that both large felids are now extirpated from NEPL, and presumably also more widely throughout Lao PDR. Mainland clouded leopard Neofelis nebulosa had the highest estimates of probability of initial occupancy, persistence and colonization, and appeared to be the most widely distributed large carnivore, followed by dhole Cuon alpinus. Both of these species emerge as a priority for further monitoring and conservation in the NEPL landscape. This study provides the most recent assessment of animal diversity and status in the NEPL. Our analytical approach provides a robust and flexible framework to include sparse and inconsistent data sets of multiple species to assess their status via occupancy as a state process, which can often provide insights into population dynamics. Keywords: Clouded leopard, Dhole, Dynamic multispecies occupancy model, Laos, Nam Et - Phou Louey National Protected Area, Tiger
... We hypothesised that (1) there is no interspecific competition among predators because tigers, leopards and dholes are able to successfully partition resources in the presence of abundant prey (Karanth and Sunquist 2000;Lovari et al. 2015), (2) tiger abundance is not influenced by single-prey availability because of their relative high abundance and variety in the study area and because, being the top predator, tigers can choose the optimal prey among the available ones (Steinmetz et al. 2013), (3) the meso-predators, leopards and dholes, are linked to suboptimal (i.e. medium-sized) prey, such as chital, nilgai or wild boar, because sambar is considered the optimal for all three predators (Karanth and Sunquist 1995;Bagchi et al. 2003b;Kumaraguru et al. 2011;Hayward et al. 2012;Mondal et al. 2012;Hayward et al. 2014) and (4) there is no interspecific competition among the prey species (i.e. there is niche segregation), because of the abundant resources in the BTR. ...
Context. Tiger (Panthera tigris), leopard (Panthera pardus) and dhole (Cuon alpinus) represent a typical multi-predator system of species of conservation concern. Several studies have addressed this system, with heterogeneous results, and there’s a lack of information on population dynamic of multi-species assemblages. We studied a time series (1998-2009) of abundance indices for three predators and five prey species in Bor Wildlife Sanctuary (BWS), Maharashtra, India, before it was declared as Bor Tiger Reserve (BTR) in 2009.
Aims. To analyse the complex relationships within a predator-prey system in a dynamic fashion: to analyse data collected in a relatively stable and undisturbed area: to form a comparison basis for future studies within the sanctuary after its declaration as a Tiger Reserve.
Methods. A 24-hr effort was made annually to census the BWS. Predators were counted at waterholes from arboreal hideouts. The prey populations were censused along 353 km line-transects. For each species we analysed yearly growth rate testing the effect of inter-species abundance.
Key results. Tiger growth rate did not depend on any particular prey, while mesopredators seemed to depend on medium-sized prey.
A die-out of dholes in 2001 was followed by an increase in tiger populations (4 to 11), which in turn negatively affected leopard numbers (6 to 2).
We found no direct evidence of top-down effect but the density dependence for three out of five prey species could be linked to predation pressure. We found some evidence of interspecific competition between prey species, especially between ungulates, potentially mediated by predation pressure.
Conclusion. The relationships between species in a predator-prey system are very complex and often could only be explained by more-than-two-species interactions. The disappearance of one predator, not necessarily the top predator, could bring multiple effects, for which it could be difficult to detect causal relationships.
Implication.
All subsequent changes in human activities in the sanctuary, as a consequence of its designation as the BTR in 2009, should be evaluated with respect to the results of this study.
The conservation of large predators should rely on the maintenance of a rich and abundant prey base in which different sized prey could lessen interactive-competition between predators.
In nutritional ecology the intake target is the diet that maximises consumer fitness. A key hypothesis of nutritional ecology is that natural selection has acted upon the behavioural and physiological traits of consumers to result in them Selectively Consuming prey to match the Intake Target (SCIT). SCIT has been documented in some herbivores and omnivores, which experience strong heterogeneity in the nutritional quality of available foods. Although carnivores experience a prey community with a much more homogeneous nutrient composition, SCIT by carnivores has nevertheless been deemed highly likely by some researchers. Here we test for SCIT for micronutrients (amino acids) in two freshwater carnivores: the river blackfish and the two-spined blackfish. Although both blackfishes exhibited non-random consumption of prey from the environment, this resulted in non-random consumption of amino acids in only one species, the river blackfish. Non-random consumption of amino acids by river blackfish was not SCIT, but instead an artefact of habitat-specific foraging. We present hypotheses to explain why wild populations of freshwater carnivores may not exhibit SCIT for amino acids. Our work highlights the need for careful, critical tests of the hypotheses and assumptions of nutritional ecology and its application to wild populations.
Cytochrome P450s are among the most important xenobiotic metabolism enzymes that catalyze the metabolism of a wide range of chemicals. Through duplication and loss events, CYPs have created their original feature of detoxification in each mammal. We performed a comprehensive genomic analysis to reveal the evolutionary features of the main xenobiotic metabolizing family: the CYP1-3 families in Carnivora. We found specific gene expansion of CYP2Cs and CYP3As in omnivorous animals, such as the brown bear, the black bear, the dog, and the badger, revealing their daily phytochemical intake as providing the causes of their evolutionary adaptation. Further phylogenetic analysis of CYP2Cs revealed Carnivora CYP2Cs were divided into CYP2C21, 2C41, and 2C23 orthologs. Additionally, CYP3As phylogeny also revealed the 3As’ evolution was completely different to that of the Caniformia and Feliformia taxa. These studies provide us with fundamental genetic and evolutionary information on CYPs in Carnivora, which is essential for the appropriate interpretation and extrapolation of pharmacokinetics or toxicokinetic data from experimental mammals to wild Carnivora.
In Southeast Asia, conservation of 'Vulnerable' clouded leopards (Neofelis nebulosa) and 'Endangered' tigers (Panthera tigris) might depend on the management of their preferred prey because large felid populations are limited by the availability of suitable prey. However, the diet of clouded leopards has never been determined, so the preferred prey of this felid remains unknown. The diet of tigers in the region has been studied only from one protected-area complex in western Thailand, but prey preferences were not determined. To better understand the primary and preferred prey of threatened felids, we used DNA-confirmed scats and prey surveys to determine the diet and prey selection of clouded leopards and tigers in a hilly evergreen forest in northern Laos. For clouded leopards, the primary prey was wild pig (Sus scrofa; 33% biomass consumed), followed by greater hog badger (Arctonyx collaris; 28%), small rodents (15%), and mainland serow (Capricornis sumatraensis; 13%; hereafter, serow). For tigers, the primary prey was wild pig (44%), followed by serow (18%), sambar (Rusa unicolor; 12%), and Asiatic black bear (Ursus thibetanus; 10%). Compared to availability, serow was positively selected by both clouded leopards (D = 0.69) and tigers (0.61), whereas all other ungulate species were consumed in proportion to the availability or avoided. Our results indicate that clouded leopards are generalist predators with a wide prey spectrum. Nonetheless, mid-sized ungulates (50-150 kg) comprised nearly half of their diet, and were the preferred prey, supporting a previous hypothesis that the enlarged gape and elongated canines of clouded leopards are adaptations for killing large prey. Because serow was the only ungulate preferred by both felids, we recommend that serow populations be monitored and managed to help conservation efforts for clouded leopards and tigers, at least in hilly evergreen forests of Southeast Asia.
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.
Habitat fragmentation is a considerable threat to biodiversity worldwide. To minimize the effects of fragmentation, it is important to identify and conserve the existing habitat connections that facilitate dispersal and gene flow among populations. Connected populations are more resilient to the changing environment that affects local populations due to greater demographic stability and higher genetic diversity. Our study is the first attempt to identify the crucial habitats facilitating the dispersal of two key sympatric cervids - spotted deer Axis axis and sambar Rusa unicolor in central India. We use species distribution models followed by landscape pattern analyses and connectivity analyses to delineate the essential habitats. Thereafter, we estimated the relative contribution of habitats outside protected areas in maintaining the ecological network, using graph-based metrics. We then locate and predict the areas that have a high risk of human-influenced cervid mortality using a Bayesian regression model that accounts for spatial structure in the data. The results show that about 55% of the core habitats, integrated across both species, lie outside the protected areas and are important in maintaining the ecological network for these cervids. Some peripheral habitats have an increased risk of anthropogenic cervid mortality, which poses high demographic risk. There is an urgent need to regulate the nature and intensity of human activities in areas of human-wildlife coexistence to maintain habitat connectivity and ensure the survival of wildlife populations. Our results on cervids complement other analyses on connectivity for large carnivores and thus enable one to account for important trophic interactions among wildlife species in land use planning.
Comprehensive information on prey availability and diet composition is important to formulate effective conservation strategies. Data on these of large apex predators, the tiger Panthera tigris, leopard Panthera pardus, and dhole Cuon alpinus, are deficient in Kalakkad-Mundanthurai Tiger Reserve, India, which harbors low prey density. Therefore, we investigated how these predators coexisted by partitioning limited food resources. We found that prey selection and activity patterns differed among predators. The leopard (D [Jacobs' preference index] = +0.916) and dhole (D = +0.937) preferred the Indian chevrotain Moschiola indica, whereas the tiger preferred the gaur Bos gaurus (D = +0.937). Biomass of tiger's diet mainly consisted of the gaur (75%). In the dhole, the Indian chevrotain and sambar Rusa unicolor together contributed 91% of biomass, whereas 79% of leopard's biomass was contributed by the Nilgiri langur Semnopithecus johnii and the Indian chevrotain. The leopard and dhole had the highest dietary overlap (Ojk [Pianka Index] = 0.46). The highest activity overlap was found among the leopard, Indian chevrotain, and sambar. The leopard and dhole would compete for the Indian chevrotain, although the competition may be avoided by different activity patterns. Managements of the Indian chevrotain and gaur are critical for conserving large predators in the region.
Context Feral cats (Felis catus), wild dogs/dingoes (Canis familiaris) and foxes (Vulpes vulpes) are predators of the endangered bridled nail-tailed wallaby (BNTW; Onychogalea frenata). Predator-proof fencing is advocated as a solution to ensure their conservation in the wild. Aims The aims of this study were to determine whether predator control translated into a reduction in their activity, find evidence of cats preying on BNTWs and understand factors that influence changes in the BNTW population size living in an unfenced reserve, particularly focusing on the influence of cat and dog control and rainfall. Methods An activity index, calculated using spoor on sand pads and images on remote cameras, was undertaken to monitor predator activity. The stomach contents of cats caught were examined to determine how commonly BNTWs feature as a prey item. The size of the BNTW population and annual survival of individuals was assessed through annual capture–mark–recapture (CMR) surveys and vehicle spotlight counts. Rainfall was measured at the study site and using data from the Bureau of Meteorology. Key results The core BNTW population estimated by CMR data increased by 214% over 4 years (2013–2017), to 400 individuals in 2017, whereas spotlight data indicated that the population had increased by 262% over 8 years (2012–2020), to 1265 individuals in 2020. The percentage of small (≤3.5 kg) BNTWs caught increased substantially over the study period. There was no detectable difference in cat or dog activity following control and no correlation was found among predator activity, rainfall and BNTW survival. The remains of BNTWs were found in 20% of cats removed from the core BNTW area. Conclusions The study confirmed that cats frequently ate BNTWs, and a combination of control methods is required to manage the threat, but there was no statistical support for a relationship between predator activity and BNTW survival. Implications The study found that native species conservation in fox-free environments is possible without the need for predator-proof fences and the ongoing maintenance costs.
Site occupancy models, accounting for imperfect detection and the influence of anthropogenic and ecological covariates, can indicate the status of species populations. They may thus be useful for exploring the suitability of landscapes such as biological corridors, to ensure population dispersal and connectivity. Using occupancy probability models of its principal prey species, we make inferences on landscape connectivity for the movement of the tiger Panthera tigris between protected areas in Bhutan. We used camera-trap data to assess the probability of site occupancy (Ψ) of the sambar Rusa unicolor, wild boar Sus scrofa and barking deer Muntiacus muntjak in biological corridor no. , which connects two national parks in central Bhutan. At least one prey species was recorded at out of trapping locations. The probability of site occupancy was highest for the barking deer (Ψ = . ± SE .) followed by sambar (Ψ = . ± SE .) and wild boar (Ψ = . ± SE .). All three species had higher occupancy probability at lower altitudes. Sambar occupancy was greater farther from settlements and on steeper and/or south-facing slopes. Barking deer also had higher occupancy on south-facing slopes, and wild boar occurred mainly close to rivers. Our findings suggest that this biological corridor could facilitate dispersal of tigers. Protecting prey species, and minimizing anthropogenic disturbance and habitat fragmentation, are vital for tiger dispersal and thus functional connectivity amongst populations in this area.
Charaspet K, Sukmasuang R, Khoewsree N, Pla-ard M, Chanachai Y. 2020. Prey species and prey selection of dholes at three different sites in Thailand. Biodiversitas 21: 5248-5262. The study of prey species and prey selection of Dholes at 3 different sites was conducted at Khao Yai National Park, Salak Pra, and Huai Kha Khaeng Wildlife Sanctuaries from 2013 to 2020. Information on Dhole prey at the sites was collected from the residues of dhole scats, from which the selection index, the relative biomass of the prey, and the relative amounts of the consumed prey were calculated. The data were collected simultaneously with the use of camera traps at each site. The study revealed that there were 13 species of Dhole prey with body weight over 5 kg. The result indicated that there were 7 species of even-toed ungulates. The relative biomass of even-toed ungulates ranged between 76.78 - 90.50% of the total biomass of all the Dholes’ consumed prey for all study sites. The dietary diversity index unveiled a similar index in all areas, which proved the adequacy of the analyzed scats. However, the Niche breadth index, which indicates the relevance of prey selection and prey species to the appearances of the prey at each site, was found to be high at Huai Kha Khaeng Wildlife Sanctuary, Khao Yai National Park, while the index was found to be low at Salak Pra Wildlife Sanctuary. The results revealed that Dholes consumed viverrid species and Malayan porcupine more often at the site where there were large carnivores. The recommendation from this study was the conservation and restoration of the ungulate populations, the main prey, as it greatly affects the conservation of the Dhole populations in Thailand. Grassland and salt lick sites, water sources improvements are also important to promote prey population. The conservation of wildlife prey by releasing them to nature, as currently conducted, has an effect on the increase of Dholes’ prey species.
Understanding traditional hunter-gatherer lifestyles in our modern world is fundamental to our understanding of their viability, as well as the role of humans as predators in structuring ecosystems. Here, we examine the factors that drive prey preferences of modern hunter-gatherer people by reviewing 85 published studies from 161 tropical, temperate and boreal sites across five continents. From these studies, we estimated Jacobs' selectivity index values (D) for 2243 species/spatiotemporal records representing 504 species from 42 vertebrate orders based on a sample size of 799,072 kill records (median = 259). Hunter-gatherers preferentially hunted 11 large-bodied, riskier species, and were capable of capturing species ranging from 0.6 to 535.3 kg, but avoided those smaller than 2.5 kg. Human prey preferences were driven by whether prey were arboreal or terrestrial, the threats the prey afforded hunters, and prey body mass. Variation in the size of prey species pursued by hunter-gatherers across each continent is a reflection of the local size spectrum of available prey, and historical or prehistorical prey depletion during the Holocene. The nature of human subsistence hunting reflects the ability to use a range of weapons and techniques to capture food, and the prey deficient wildlands where people living traditional lifestyles persist.
Inferring the mechanisms that facilitate sympatry amongst large mammalian carni-vores in anthropogenically altered habitats is critical if conservation initiatives aimed to recover populations are to be effective. In this study, we assessed the utilization of space, time and prey resources by tigers Panthera tigris, leopards P. pardus and dholes Cuon alpinus in Manas National Park (MNP), India-a site where prey densities are not limiting, yet anthropogenic influences alter the spatial and temporal behaviour of prey. We use photographic capture data and predator scats collected over three sampling years (2014-15, 2015-16 and 2016-17) to assess patterns of (a) spatial use using a multispecies species occupancy framework , (b) time-activity patterns and overlap between predators and prey using non-parametric circular kernel-density functions, (c) fine-scale spatio-temporal behaviour by comparing time-to-encounters of subsequent events and (d) predator diets by analysing prey remains in predator scats. Our results highlight that the predators segregate through fine-scale spatio-temporal avoidance rather than displaying population level changes in space-use, activity patterns or food habits. Overlap in space-use between tigers and leopards was high and time-activity patterns of the predators closely matched those of prey, suggesting that predators likely maximize resource acquisition in this prey-rich environment. Ungulate prey dominated the diet of predators, resulting in the high dietary overlap. From our results, we infer patterns of sympatry among large carnivores in the face of anthropogenic influences and highlight the need to understand interspecific interactions within a community before initiating conservation actions aimed at recovering these endangered species.
The Asian golden cat (Catopuma temminckii) occurs in small, declining, and highly fragmented populations throughout Southeast Asia, whereas the smaller leopard cat (Prionailurus bengalensis) is common and widespread. In contrast to leopard cats, little is known about the ecology of Asian golden cats, and resource partitioning between these species has not been studied. We used DNA-confirmed scats, camera-trap data, and prey surveys, to determine the diet, prey selection, and activity, of Asian golden cats and leopard cats in a protected area in northern Laos. The two felids had different diets: Asian golden cats consumed mostly ungulates (35% biomass consumed), murid rodents (23%), and carnivores (15%), whereas leopard cats consumed mostly murid rodents (79%). Asian golden cats were not random in their consumption of ungulates, because muntjac (Muntiacus spp.) were selectively consumed over larger ungulates, indicating muntjac were preyed upon rather than scavenged. Dietary overlap between the two felid species was moderate (R 0 = 0.60), and the dietary niche breadth of Asian golden cats (B = 8.44) was nearly twice as high as that of leopard cats (4.54). The mean (± SD) scat diameter was greater for Asian golden cats (2.1 ± 0.3 cm) than leopard cats (1.8 ± 0.2 cm), although diameters of leopard cat scats were considerably larger than previously assumed for this species. The felid species differed in their activity patterns, because Asian golden cats were diurnal, whereas leopard cats were nocturnal, although they did not differ in their use of elevation, suggesting there was no habitat segregation. Overall, leopard cats appeared to coexist with Asian golden cats, a potential predator and competitor, by exhibiting dietary and temporal partitioning. Our results showed that muntjac were important prey of Asian golden cats, suggesting the management of muntjac might be important for conserving populations of Asian golden cats.
Longitudinal studies of wildlife are urgently needed in South‐East Asia to understand population responses to the high poaching pressure that characterizes this region. We monitored population trends and habitat use of five heavily poached ungulate species (gaur, sambar, wild pig, red muntjac, and Fea's muntjac) over five years in two protected areas in western Thailand using camera trap surveys. We used single‐season occupancy models to investigate effects of ecological and anthropological variables on ungulate distribution, and multi‐season models to assess occupancy dynamics over time. Occupancy of gaur and sambar was low (<0.25), but concentrated near saltlicks and at low elevations. Wild pig and muntjac occupancies were 3–4 times higher (0.60–0.80). Wild pig occupancy was lower near villages, but this effect dissipated in the final year of the study, coinciding with a purported decrease in poaching. Wild pig occupancy increased significantly, with the probability of colonizing new sites doubling from 0.40 to 0.81 over time. In contrast, occupancy rates of gaur, sambar, and muntjac did not grow, though they were stable. Poaching pressure during the study was low, perhaps allowing populations to stabilize. But only wild pig (the most resilient of the five species) increased. The failure of gaur and sambar to recover might stem from historical overhunting combined with ecological constraints, particularly low saltlick density. Recovery of ungulates (and the carnivores that depend on them) in overhunted South‐East Asian reserves might require intensive interventions, particularly habitat improvement and population augmentation, to achieve conservation objectives.
The total number of white blood cells (WBCs) is related the immune system. In mammals, it is affected by the body mass, but it is unclear how the numbers of different WBC types correlate with this parameter. We analyzed the effect of body mass on WBC number and ratio in felids, where species are similar in diet (warm-blood vertebrates) and reproductive strategy (promiscuity). Based on zoo veterinary data (ZIMS database) we analyzed the effect of body mass on WBC number and neutrophils/lymphocytes ratio in 26 species of felids. The number of WBCs correlated with the body masses of animals: large cats had more WBC, which may be due to greater risks of infection associated with larger body surface, lifespan and home range size. For the first time we found obvious differences in the number of WBC types. Large cats also had more neutrophils and monocytes but fewer lymphocytes than smaller cats. The ratio of neutrophils to lymphocytes is greater in large felids. This phenomenon may be related to diet (relative prey size and kill utilization time), which suggests regular contact of large cats with bacterial and protozoal pathogens in contrast to the small cats.
Striped hyenas (Hyaena hyaena) are extremely rare in Nepal, and only a few people have studied them in their natural forest and grassland habitat. Their rarity is due to anthropogenic pressures such as hunting, habitat modification, being killed on roads, and depletion of their natural prey. Here, we studied the feeding ecology of hyenas in lowland, Nepal. We employed an opportunistic sampling to collect hyena scats in a range of habitats and the line transect sampling to identify the prey of the hyena in the study site. We collected 68 hyena scats between 2015 and 2018. Most of the hyena scat (39.7%) was found in the Churia Hill forest followed by riverbed (26.4%), mixed forest (14.7%), Sal (Shorea robusta)‐dominated forest (11.7%), and grassland area (7.3%). We found eleven mammalian prey species, plants, and some unidentified items in the hyena scats. The frequency of occurrence and relative biomass of the medium‐sized wild boar (Sus scrofa) were higher than other smaller prey species such as hare (Lepus nigricollis) and rhesus macaque (Macaca mulatta). Similarly, the proportion of large prey species such as nilgai (Boselaphus tragocamelus) in the hyena diet was lower compared with wild boar, hares, and rhesus macaques indicating medium‐sized wild boar is the most preferred prey species. Livestock contributed 17.3% of the total dietary biomass. Domesticated species such as goats, sheep, cows, and even dogs were found in the diet of hyenas. Predation of livestock by hyenas could cause conflict, especially if this ongoing issue continues in the future. Rather, more conservation effort is required in lowland areas of Nepal to protect the hyenas' natural prey species, particularly in wildlife habitats to reduce the lure of taking domestic livestock. Similarly, conservation education at the local level and active involvement of government authorities in the conservation of this species might be helpful to mitigate human–hyena conflict in the human‐dominated landscape.
Large carnivores have experienced significant global range contractions and recovering their populations is often contingent on increasing prey abundances. In Manas National Park (MNP), following nearly two decades of ethnopolitical conflict, studies document that populations of both prey and predators were depressed. Here we assess the status of wild ungulates in a section of MNP (Bansbari-Bhuyanpara) that has remained conflict-free for over a decade. For seven ungulate species, we estimate species-specific densities using distance-based sampling, assess species-specific space-use patterns in relation to habitat variables within an occupancy framework and examine patterns of temporal activity in relation to times when people access the park for resources. Further, by comparing temporal activity patterns of ungulates between MNP, a site where local communities access the park for resources, and Kaziranga National Park, where human use of the park is minimal, we examine if species activity is altered in response to human presence. We estimate that currently Bansbari-Bhuyanpara ranges of MNP support 42.66 (34.16-51.16) individual ungulates/km 2. Our results highlight that current patterns of human access within the park affect both spatial and temporal behaviour of these species. Although we estimate a relatively high recovery potential for tigers in MNP given current prey densities, we suggest that further ungulate population recoveries could be supported in the park. With several ungulate species experiencing range-wide declines, efforts to minimize non-lethal human disturbances on these species also need to be considered to ensure that predator-prey systems remain intact.
Dramatic population declines threaten the Endangered Indochinese tiger Panthera tigris corbetti with extinction. Thailand now plays a critical role in its conservation, as there are few known breeding populations in other range countries. Thailand's Dong Phayayen-Khao Yai Forest Complex is recognized as an important tiger recovery site, but it remains poorly studied. Here, we present results from the first camera-trap study focused on tigers and implemented across all protected areas in this landscape. Our goal was to assess tiger and prey populations across the five protected areas of this forest complex, reviewing discernible patterns in rates of detection. We conducted camera-trap surveys opportunistically during 2008–2017. We recorded 1,726 detections of tigers in 79,909 camera-trap nights. Among these were at least 16 adults and six cubs/juveniles from four breeding females. Detection rates of both tigers and potential prey species varied considerably between protected areas over the study period. Our findings suggest heterogeneity in tiger distribution across this relatively continuous landscape, potentially influenced by distribution of key prey species. This study indicates that the Dong Phayayen-Khao Yai Forest Complex is one of the few remaining breeding locations of the Indochinese tiger. Despite limitations posed by our study design, our findings have catalysed increased research and conservation interest in this globally important population at a critical time for tiger conservation in South-east Asia.
Felids show remarkable phenotypic similarities and are conservative in behavioral and ecological traits. In contrast, they display a large range in body mass from around 1kg to more than 300kg. Body size and locomotory specializations correlate to skull, limb and vertebral skeleton morphology. With an increase in body mass, felids prey selection switches from small to large, from using a rapid skull or spine lethal bite for small prey, to sustained suffocating bite for large prey. Dietary specialization correlates to skull and front limbs morphology but no correlation was found on the spine or on the hind limb. The morphology of the sacroiliac junction in relation to ecological factors remained to be described. We are presenting a study of the overall shape of the iliac auricular surface with qualitative and quantitative analyses of its morphology. Our results demonstrate that body mass, prey selection, and bite type, crucially influence the auricular surface, where no significant effect of locomotor specialization was found. The outline of the surface is significantly more elevated dorso-caudally and the joint surface shows an irregular W-shape topography in big cats whereas the surface in small cats is smoother with a C-shape topography and less of an elevated ridge. Biomechanically, we suggest that a complex auricular surface increases joint stiffness and provides more support in heavier cats, an advantage for subduing big prey successfully during a sustained bite.
Habitat loss, prey depletion, and direct poaching for the illegal wildlife trade are endangering large carnivores across the globe. Tigers (Panthera tigris) have lost 93% of their historical range and are experiencing rapid population declines. A dominant paradigm of current tiger conservation focuses on conservation of 6% of the presently occupied tiger habitat deemed to be tiger source sites. In Bhutan, little was known about tiger distribution or abundance during the time of such classification, and no part of the country was included in the so-called 6% solution. Here we evaluate whether Bhutan is a potential tiger source site by rigorously estimating tiger density and spatial distribution across the country. We used large scale remote-camera trapping across n = 1129 sites in 2014–2015 to survey all potential tiger range in Bhutan. We estimated 90 individual tigers (60 females) and a mean density of 0.23 adult tigers per 100 km2. Bhutan has significantly higher numbers of tigers than almost all identified source sites in the 6% solution. With low human density and large swaths of forest cover, the landscape of Bhutan and adjacent northeast India is a promising stronghold for tigers and should be prioritized in large-scale conservation efforts.
Home range and food habits of tigers (Panthera tigris tigris) were studied in Sariska Tiger Reserve from July 2008 to June 2009. Three tigers (one male and two females) were radio-collared and reintroduced in Sariska Tiger Reserve from Ranthambhore Tiger Reserve, Western India during 2008-2009. The reintroduced tigers were monitored periodically through ground tracking using "triangulation and homing in techniques." The estimated annual home ranges were 168.6 km2 and 181.4 km2 for tiger and tigress-1 respectively. The estimated summer home range of tigress-2 was 223.4 km2. In total, 115 kills and 103 scats of tigers were collected to study the food habits. The line transect method was used to estimate the prey availability. The density of peafowl (Pavo cristatus) was found to be highest (125.2 ± 15.3/ km2) in Sariska followed by livestock (Bubalis bubalis and Bos indicus) (59.9 ± 22.3/ km2), chital (Axis axis) (46.7 ± 9.5/ km2), sambar (Rusa unicolor) (26.2 ± 4.9/ km2), common langur (Semnopithecus entellus) (22.8 ± 6.5/ km2), nilgai (Boselaphus tragocamelus) (19.5 ± 3.3/ km2) and wild pig (Sus scrofa) (15.4 ± 4.4/ km2). Tigers fed on seven prey species as shown by kill data. Tigers' scat analysis revealed the presence of five prey species. Prey selection by tigers based on scat analysis was in the following order: sambar> chital> nilgai> livestock> common langur. It is proposed to restock the tiger population initially with five tigers in Sariska and subsequent supplementation of two tigers every three years for a period of six years, which will allow the population to achieve demographic viability. Removal of anthropogenic pressure from the national park will be crucial for the long term survival of tigers in Sariska. © K. Sankar, Qamar Qureshi, Parag Nigam, P.K. Malik, P.R. Sinha, R.N. Mehrotra, Rajesh Gopal, Subhadeep Bhattacharjee, Krishnendu Mondal and Shilpi Gupta.
Ecological factors influencing prey selection and prey removal by tigers (Panthera tigris) during the dry season were investigated in the Karnali floodplain in the southwestern part of Royal Bardia National Park. Nepal, between 1990 and 1994. Densities of large herbivore prey were estimated by line transects. Density of tigers was estimated from individual track features. Tiger diet was determined by analyses of scats (feces). Selection of prey species was inferred by comparing observed counts of scats to hypothesized scat frequencies generated from prey density estimates. Prey removal by tigers was estimated from the estimated food requirement of the tigers in the study area and the estimated biomass of the different prey species. Three species (axis deer (Axis axis), wild boar (Sus scrofa) and hog deer (Axis porcinus)) comprised more than 90% of the biomass killed. Larger and reportedly more optimum sized prey species like nilgai (Boselaphus tragocamelus) and barasingha (Cervus duvauceli) were not preferentially eaten, probably due to their low densities. Contrary to the predictions that tiger predation would be non-selective on medium and smaller sized prey if availability of large prey was limited, the Karnali tigers showed significant selectivity among the medium sized prey species: wild boar was preferred, whereas barking deer (Muntiacus muntjak) was under-represented in the diet. Total removal of available ungulate prey biomass was low (3.1%) compared to other studies, indicating that the relatively dense tiger population was regulated through social behaviour rather than by food. Predation by tiger did not appear to play a major role in regulating the densities of wild herbivores, except nilgai. In spite of high biomass and diversity of prey, leopard (Panthera pardus) was seemingly displaced to the periphery of the park, suggesting that interspecific predation rather than food competition excluded coexistence with tigers.
Lynx Lynx lynx (Linnaeus, 1758) - ungulate relationships were studied in the exploited (530 km2) and pristine (47.5 km2) parts of Bialowieza Primeval Forest during 1985 through 1992. In pristine forest, Sus scrofa (54%) and Cervus elaphus (35%) predominated in the ungulate community. In exploited forest, C. elaphus (36%) and Capreolus capreolus (33%) were most abundant.
Despite a [greater-than-or-equal]75% reduction in the geographic range of Mongolian gazelles Procapragutturosa over the past 50 years, the species is still recognized as the most numerous large grassland herbivore in Asia. Its actual population size, however, is still disputed, and we therefore estimated its numbers in an 80,000 km2 area in the eastern steppe of Mongolia by driving long-distance (1,2002002. Quantitative estimates of gazelle numbers are essential for understanding the causes of changes in the population, and thus devising conservation strategies to assure its long-term viability. Observed herds ranged in size from 142. Density estimates varied from 10.7 gazelles km2 in autumn, with total population estimates of 803,820 (483,7901,491,278 95% confidence interval), respectively. Confidence limits were wide, and to obtain a coefficient of variation of 20%, transect lengths would need to be extended three- to four-fold. Until more efficient means for conducting population surveys can be implemented, driving long-distance transects, combined with distance analysis, seem to provide the best quantitative estimate of Mongolian gazelle populations.
Synopsis The tiger (Panthera tigris) is the largest obligate predator in the Asian temperate and tropical forest ecosystems in which it occurs. The plasticity and constraints in tiger resource acquisition are examined in the context of abrupt and pervasive environmental change throughout the tiger's rangechanges that threaten its survival. Prey capture in tigers is plastic, allowing tigers to capture prey of a wide range of sizes and types. In Nepal's Chitwan National Park, tigers selected large cervids, thereby gaining access to a substantial proportion of the ungulate biomass which is not available to smaller felids. In mainland environments where primary prey are larger ungulates, tigers are larger, as much as twice the size of the Sunda Island tigers. Large body size may increase the efficiency of preying on big ungulates, but decreases the efficiency of living on smaller prey types. The smaller body size of the Sunda Island tigers may increase the efficiency of capturing the smaller prey that are relatively abundant in rainforest environments, feeding primarily on plant reproductive parts, but does not preclude the capture of large prey, such as Bos (800 kg). Understanding tiger resource acquisition is important to tiger conservation. The integrity of tiger ecotypes should be recognized in the management of captive and wild tigers as a metapopulation or metapopulations. The loss of larger prey types from habitat fragments can be expected to severely affect the survival of tigers in those habitats; larger tigers should be more severely affected than smaller tigers because of their different success-of-capture curves. The extirpation of tigers from an area can result in changes in the relative abundance of large ungulates and may result in an increase in the number of smaller predators, thus altering community structure in these ecological systems. Tigers do not kill human beings in numbers proportionate to their availability and their potential vulnerability. Killing of humans becomes a problem when individual tigers are excluded from normal prey populations through social processes and/or
Niches of 2 cervids (chital, Axis axis and sambar, Cervus unicolor) and 2 bovids (nilgai, Boselaphus tragocamelus and chinkara, Gazella bennetti) from semiarid forests in western India were studied for habitat use and food habits. Habitat use was analyzed by discriminant analysis using 20 variables, and food habits investigated by analyzing undigested plant remains in pellets. Cervids and bovids differentiated primarily according to vegetation and terrain features, and the 2 deer species showed separation in diet. The 2 cervids selected forested areas, whereas the 2 bovids selected scrubland and were more tolerant of disturbances like livestock grazing and also showed a high similarity in food habits. Habitat use and food habits were analyzed with nonmetric multidimensional scaling to assess their combined effects. A greater degree of similarity in resource use between the 2 bovids suggests that they may be competitive, at least during periods of forage scarcity.
The study provides information on distribution, abundance of wild ungulates and their link to habitat characteristics at a landscape level conservation. Khata Corridor is an important component of Terai Arc Landscape. It is about 9 km long and connects Bardia National Park, Nepal with Katarniaghat Wildlife Sanctuary, India. Relative abundance and distribution of Tiger prey base species (wild ungulates) was studied from mid April to mid June 2008 at Khata Corridor. A total of 127 transects (1 transect=625m, distance between adjacent transect=100m, distance between two parallel transect=100m) containing 2700 circular plots of 10m2 were laid and sampled following Smith et al (1999). A total of 2043 wild ungulates pellets were counted within the sampled area with a mean pellet group abundance of 0.75 pellets per plot. Mean pellet group abundance of Spotted deer, Wild boar, Monkey, Hog deer, Barking deer and Blue bull were 0.63, 0.06, 0.05, 0.02, 0.00 and 0.00 per plot respectively. Species wise ungulate abundance showed that Barking deer was highest in Low Density Mixed Forest, Spotted deer in Grassland, Hog deer in both Grassland and Low Density Mixed Forest, Blue bull only in Low Density Mixed Forest, Monkey in Riverine Forest and Wooded Grassland, and Wild boar in Wooded Grassland. Distribution pattern of wild ungulate species was of Clumped type (S2/a=93.10). Out of four habitat types, flood plains with riverine forest and grasslands were considered important habitats. Khata has been experiencing different levels of degradation also has tremendous conservation potential to ensure long term viability of large mammals (eg. Tigers) if restored and integrated into landscape level conservation plans.
Recent technological advances in Global Positioning system (GPS) technology have
generated an increase in the use of GPS collars fitted to wild animals to track their move-
ments. GPS units are advantageous compared to other methods of remote tracking of
animals in that the have the ability to gather large quantities of spatial data that may be
accessed remotely. However, animal behaviour, topography, vegetation, and seasonality
could all influencevGPS acquisition success and therefore bias results from GPS technology.
Here we analyse temporal and seasonal patterns associated withGPSacquisition failures in
five GPS collars fitted to leopards (Panthera pardus) in the Waterberg region of South Africa.
Of 6565 attempted GPS acquisitions, 19% (1236) failed with the location failure rate of
individual collars ranging from 15–29%. Single failed attempts accounted for the majority of
location failures, suggesting that failure was caused by temporary blockage to satellites.
Failure rates were significantly higher during the day and there were indications that they
were higher during the wet season than in the dry season. Failures were also clustered in
space for some individual animals. Our results suggest that temporal patterns of animal
behaviour and habitat choice influence the probability of GPS location failures in this
species. We therefore suggest that potential biases should be accounted for when using
GPS data to analyse movement and habitat selection in this and similar carnivore species.
Knowledge of the range, behavior, and feeding habits of large carnivores is fundamental to their successful conservation.
Traditionally, the best method to obtain feeding data is through continuous observation, which is not always feasible. Reliable automated
methods are needed to obtain sample sizes sufficient for statistical inference. Identification of large carnivore kill sites using Global Positioning
System (GPS) data is gaining popularity. We assessed performance of generalized linear regression models (GLM) versus classification trees
(CT) in a multipredator, multiprey African savanna ecosystem. We applied GLMs and CTs to various combinations of distance-traveled data,
cluster durations, and environmental factors to predict occurrence of 234 female African lion (Panthera leo) kill sites from 1,477 investigated
GPS clusters. Ratio of distance moved 24 hours before versus 24 hours after a cluster was the most important predictor variable in both GLM
and CT analysis. In all cases, GLMs outperformed our cost-complexity-pruned CTs in their discriminative ability to separate kill from nonkill
sites. Generalized linear models provided a good framework for kill-site identification that incorporates a hierarchal ordering of cluster
investigation and measures to assess trade-offs between classification accuracy and time constraints. Implementation of GLMs within an
adaptive sampling framework can considerably increase efficiency of locating kill sites, providing a cost-effective method for increasing sample
sizes of kill data.
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.
Data on lion home range size, pride size, density, group size, cub survival, litter size and adult sex ratio were extracted from studies carried out in 10 habitats. The data were compared with five measures of food supply: mean prey biomass, prey biomass during the season of greatest abundance, prey biomass during the season of least abundance, biomass of middle-sized prey species (i.e. those with mean adult live-weights between 50 and 250 kg), and the mean size of carcasses fed on by lions. The results indicated that range size is inversely correlated with the abundance of prey during the period of least abundance, but not with overall prey abundance. Range size showed no consistent variation with pride size or with minimum metabolic requirements. Pride size (measured in terms of average number of animals per pride and average number of adult females) and cub survival strongly correlated with lean season food abundance. No relationship was found between group size or litter size and any of the measures of food supply Adult sex ratio did not vary consistently with food supply or lion density, although the data did suggest that prides inhabiting small, circumscribed reserves may experience less inter-male competition and this, in turn, may affect the adult sex ratio.
The mammalian fauna of Africa, and particularly its ungulate component, has evolved to a variety and multitude unparalleled elsewhere—at least in recent times—both in terms of different species and of sheer numbers. Its diversity and profusion have greatly impressed Western man ever since he began invading and colonizing Africa, and continue to do so to the present day. Until quite recently, large tracts of land, with their natural habitats and their wild animals, have remained relatively unaffected by “civilization”, but the latter is advancing inexorably in all directions, and truly “wild places” become ever harder to find. Yet, at the same time, Western man experiences a growing need for just such places, to escape from his overpopulated and polluted surroundings and to re-establish some long-lost contact with “nature.” The recent upsurge of tourism in many African countries bears witness to this need, and the African ungulates contribute a large share in attracting this tourism, which provides substantial earnings for the host countries.
This chapter provides new insights into the tiger's ability and flexibility to persist in current and changing landscapes. These insights come from a variety of sources, including long-term telemetry studies of tigers in southern Nepal and the Russian Far East, sites with vastly different ecological conditions, as well as from long-term monitoring of tiger and prey populations in Nagarahole National Park, India, and from dietary studies and prey density estimates at other sites in India. The development of camera-trapping techniques has also provided the first reliable density estimates of tiger populations in the lowland tropical rainforests of Thailand, Sumatra, Peninsular Malaysia, and Laos. This chapter synthesizes these studies to describe tiger behavior and ecology. It describes the tiger's basic skill set, and then reviews predatory behavior and ecology, followed by population ecology. Finally, this chapter concludes with some thoughts about research directions and development of new methods of enumerating prey density and assessing tiger diets in these habitats.
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.
1. Ecological factors influencing prey selection by tiger Panthera tigris, leopard Panthera pardus and dhole Cuon alpinus were investigated in an intact assemblage of large mammals in the tropical forests of Nagarahole, southern India, between 1986 and 1990. 2. Densities of large herbivores were estimated using line transects, and population structures from area counts. Carnivore diets were determined from analyses of scats (faeces) and kills. Selectivity for prey species was inferred from likelihood ratio tests comparing observed counts of scats to hypothesized scat frequencies generated from prey density estimates using parametric bootstrap simulations. Predator selectivity for size, age, sex and physical condition of prey was estimated using selection indices. 3. Ungulate and primate prey attained a density of 91 animals km-2 and comprised 89-98% of the biomass killed. Predators showed significant (P
Long-term continuous observations of hunting lions Panthera leo in the Kruger National Park were used to assess the variables affecting hunting success of male and female lions. Generalized linear models revealed that seven variables had significant independent influences on hunting success, with the most important being the prey species hunted. Three types of variables were recognized: (1) lion related, where type of hunt, wind orientation, and the number of adults hunting; (2) prey related, where prey species and herd size; (3) environment related, where moon brightness, and grass height were significant. The sex of the lions had no effect on the overall probability of hunting success. Five second-order interactions significantly influenced the probability of hunting success, with the most important being the interaction between sex and the type of prey. The only significant third-order interaction containing the variable sex, was the inter-relationship with prey species and shrub cover. After removing the over-riding bias of the prey species, greater resolution of the factors affecting success was revealed. The major difference was that group size influences hunting success and concomitantly prey selection, promoting selection for medium-sized ungulates like zebra Equus burchelli and wildebeest Connochaetes taurinus by females, and males mainly capturing buffalo Syncerus caffer. We conclude that in African ecosystems, the hunting success of male and female lions varies with a range of combinations of lion-, prey- and environment-related variables. We demonstrate the hunting ability of male lions, which has perhaps been understated in other studies.
This paper examines the hypothesis that in the field redshank select the sizes of polychaete worms which maximize the biomass ingested per unit time (ingestion rate). Feeding rate (number taken per metre) on large worms depended mainly on their density in the mud. Feeding rate on small worms was more influenced by the biomass ingested from large worms than by their own density so that small worms were only taken frequently when large ones were scarce, suggesting that small prey were least preferred. Experiments with a mathematical model of the birds foraging behaviour suggested that the tendency to vary their responsiveness to small worms as a function of the biomass ingested from large ones did indeed maximize ingestion rate.
The majority of individuals in a community belong to a small number of abundant species. Understanding why some species are rare and others are common has been a long-held goal for ecologists. Africa’s large carnivore guild preferentially preys on a small number of species within a limited weight range. Within this weight range however, some species that are expected to be significantly preferred as prey are not. I tested whether these species avoid preferential predation through their low densities. Records of over 40,000 kills from up to 48 different communities were used to test if non-preferred species within the expected prey weight ranges of each large predator avoid preferential predation and why. Species expected to be prey of Africa’s large predators based on their body mass, that are preferred are preyed upon significantly more frequently at low densities than non-preferred prey. This results in a negative relationship between relative abundance and preference for preferred prey, but a positive relationship for non-preferred prey. The non-preferred prey species that are within the expected prey weight ranges of Africa’s large predators are significantly less abundant within the prey community than significantly preferred prey. Rarity in African ungulates may convey an anti-predator benefit in that it was suboptimal for predators to evolve morphological or behavioral strategies to optimally forage on them or in that prey species can avoid predators by existing in habitats with low carrying capacity.
The primary factor which facilitated prey resource partitioning among dhole (Cuon alpinus), leopard (Panthera pardus) and tiger (P. tigris) in Bandipur Tiger Reserve, South India was their size which in turn determined the size of the prey killed. Kills over 100 kg formed 42 % of the tiger kills (n = 19) ; 69 % of the kills by leopard weighed less than 50 kg (n = 58) as did 74 % of dhole kills (n = 302). Dholes, the coursing predator, killed more chital (Axis axis), and sambar (Cervus unicolor) males and there seems to be a trend for tiger and leopard (stalkers) killing more does. The data suggest that prime age chital were relatively more free from dhole predation than the past prime animals. Dholes killed more chital fawns when compared with leopard kills. Chital, by its behaviour of yarding in open areas at night in and around Bandipur village in large groups, avoided tiger predation while sambar by preferring dense areas and living in smaller groups made itself more vulnerable. To understand the precise interactions among these predators and with their prey species further research in habitats where coursers and stalkers co-exist is suggested.