Morten Odden

Big Cats in Our Backyards: Persistence of Large Carnivores in a Human Dominated Landscape in India

Protected areas are extremely important for the long term viability of biodiversity in a densely populated country like India where land is a scarce resource. However, protected areas cover only 5% of the land area in India and in the case of large carnivores that range widely, human use landscapes will function as important habitats required for gene flow to occur between protected areas. In this study, we used photographic capture recapture analysis to assess the density of large carnivores in a human-dominated agricultural landscape with density >300 people/km2 in western Maharashtra, India. We found evidence of a wide suite of wild carnivores inhabiting a cropland landscape devoid of wilderness and wild herbivore prey. Furthermore, the large carnivores; leopard (Panthera pardus) and striped hyaena (Hyaena hyaena) occurred at relatively high density of 4.8±1.2 (sd) adults/100 km2 and 5.03±1.3 (sd) adults/100 km2 respectively. This situation has never been reported before where 10 large carnivores/100 km2 are sharing space with dense human populations in a completely modified landscape. Human attacks by leopards were rare despite a potentially volatile situation considering that the leopard has been involved in serious conflict, including human deaths in adjoining areas. The results of our work push the frontiers of our understanding of the adaptability of both, humans and wildlife to each other’s presence. The results also highlight the urgent need to shift from a PA centric to a landscape level conservation approach, where issues are more complex, and the potential for conflict is also very high. It also highlights the need for a serious rethink of conservation policy, law and practice where the current management focus is restricted to wildlife inside Protected Areas.

Translocation as a Tool for Mitigating Conflict with Leopards in Human-Dominated Landscapes of India

We examined the efficacy of a translocation program in which large numbers of leopards (Panthera pardus fusca) were trapped in human-dominated landscapes where livestock attacks were common and human attacks rare and released into adjoining forested areas in an attempt to reduce leopard presence and mitigate conflicts at the capture site. In the year starting in February of 2001, 29 leopards were captured in the human-dominated rural landscape of the Junnar region (4275 km(2) , 185 people/km(2) ), Maharashtra, India, and released an average of 39.5 km away in adjoining forests. Eleven leopards were also relocated to the same forests from other districts. Prior to the large-scale translocation program, an average of four leopard attacks on humans occurred each year between 1993 and 2001. After the translocation program was initiated, the average increased substantially to 17 attacks. Linear and logistic models showed that attack frequency increased in Junnar following nearby releases of leopards and decreased when leopards were removed for releases far away; that attacks became more lethal when the number of leopards introduced from other districts increased; and that attacks were most likely to occur in the regions where the largest number of leopards had been introduced from other areas. These results suggest that leopards did not stay at the release sites and that translocation induced attacks on people. Potential explanations for these results include increased aggression induced by stress of the translocation process, movement through unfamiliar human-dominated landscapes following release, and loss of fear of humans due to familiarity with humans acquired during captivity. Our results show that reactive solutions to attacks on humans by leopards, such as translocation, could in fact increase human-leopard conflict. Measures to reduce human-carnivore conflicts may include more effective compensation procedures to pay livestock owners for the loss of animals to predation by carnivores, providing better methods of protection for livestock, and encouraging greater social acceptance of the presence of carnivores in human-dominated landscapes.

Habitat and roe deer fawn vulnerability to red fox predation

1. Notwithstanding the growing amount of literature emphasizing the link between habitat, life-history traits and behaviour, few empirical studies investigated the combined effect of these parameters on individual predation risk. We investigated direct and indirect consequences of habitat composition at multiple spatial scales on predation risk by red foxes on 151 radio-monitored roe deer fawns. We hypothesized that the higher resource availability in fragmented agricultural areas increased predation risk because of: (i) shorter prey movements, which may increase predictability; (ii) larger litter size and faster growth rates, which may increase detectability in species adopting a hiding neonatal anti-predator strategy. The sharing of risky habitat among littermates was expected to promote whole-litter losses as a result of predation. 2. The landscape-scale availability of agricultural areas negatively affected pre-weaning movements, but did not influence growth rates or litter size. Predation risk was best described by the interplay between movements and fine-scale habitat fragmentation: a higher mobility increased the encounter rate and predation risk in highly fragmented home ranges, while it reduced predation risk in forest-dominated areas with clumped resources because of decreased predictability. This is one of the first demonstrations that movement patterns can be an efficient anti-predator strategy when adjusted to local conditions. 3. In accordance with previous studies documenting the existence of family effects (i.e. non-independence among siblings) in survival, littermates survived or died together more often than expected by chance. In addition, our study specifically demonstrated the occurrence of behaviourally mediated family effects in predation risk: after a fox killed one fawn the probability of a sibling being killed within a few days rose from 20% to 47%, likely because of the win-stay strategy (i.e. return to a previously rewarding site) adopted by the predator. Hence, the predators hunting strategy has the potential to raise fawn mortality disproportionately to predator abundance. 4. There is increasing evidence that fawns inhabiting highly productive predator-free habitats are granted lifetime fitness benefits; these potential advantages, however, can be cancelled out when predation risk increases in the very same high-productivity areas, which might thus turn into attractive sinks.

Do tigers displace leopards? If so, why?

We investigated predictions concerning the competitive relationships between tigers Panthera tigris and leopards Panthera pardus in Bardia National Park, Nepal, based on spatial distributions of scats and territorial markings (sign), analyses of scat content and census of wild ungulate prey. Medium-sized ungulates, in particular chital Axis axis, was the main food of both predators, but leopards consumed significantly larger proportions of domestic animals, small mammals, and birds than tigers. Tiger sign were never found outside the park, while leopard sign occurred both inside and outside, and were significantly closer to the park border than tiger sign. Significantly higher prey densities at locations of tiger sign than that of leopards were mainly due to a preference of the latter species for the park border areas. Our results imply that interference competition––and not competition for food––was a limiting factor for the leopard population, whose distribution was restricted to the margins of the tiger territories. We suggest that the composition of the prey base is a key factor in understanding the different results and interpretations reported in studies on tiger/leopard coexistence. There are two potential mechanisms that link interference competition and prey: (1) low abundance of large ungulate prey decreases foraging efficiency of tigers, leading to increased energetic stress and aggression towards leopards; and (2) increased diet overlap due to scarcity of large prey leads to increased encounter rates and increased levels of interference competition.

When a generalist becomes a specialist: patterns of red fox predation on roe deer fawns under contrasting conditions

The red fox (Vulpes vulpes (L., 1758)) functional response to roe deer (Capreolus capreolus (L., 1758) fawn density was investigated in two Norwegian study areas characterized by a 26-fold differen...

Spacing and activity patterns of leopards Panthera pardus in the Royal Bardia National Park, Nepal

Abstract Space use and activity of radio-collared leopards Panthera pardus (two adult males and one adult female) were monitored during 3–25 months in a prey-rich part of the Royal Bardia National Park, Nepal. Annual home ranges of the two males were 47 and 48 km2 and had an overlap of only 7%, whereas the overlap between the females home range (17 km2) and that of one of the males was 56%. The range sizes were larger than reported from other studies in southeast Asia, but much smaller than some ranges in Africa. When comparing different studies, the sexual difference in range size increased significantly with increasing average range size. Thus, the cost by males of traversing large home ranges is probably not a determinant factor in shaping leopard communities. The females seasonal home ranges (5.2 and 6.6 km2) were smallest during the seasons when her cubs were less than six months of age. She moved her home range closer to agricultural fields during the season when the abundant and important prey axis deer Axis axis visits these areas most frequently. No such pattern was detected among the two males. Instead they frequented human settlements throughout the year, probably in order to hunt easily accessible domestic animals. Home ranges in similar seasons in consecutive years overlapped more (female = 64%, male = 75%) than ranges in different seasons in the same year (female = 38%, male = 64%). Intensive tracking sessions of 24 hours revealed that the diel activity levels of the two sexes were similar (female: 62.3%, males: 62.6%). However, their patterns of activity were different as the males moved mainly at night (day: 1,582 m, night: 5,244 m) and the female moved similar distances day and night (day: 2,381 m, night: 2,698 m). The female may have restricted her movement at the time when conspecific males were likely to be active in order to protect her cubs from infanticide. The males moved in a more linear manner than the female, and the linear distances between radio locations from consecutive days differed significantly between the sexes (male: 3,324 m, female: 881 m), but the actual distances moved during the 24-hour cycles were fairly similar (male: 6,826 m, female: 5,079 m).

A cat among the dogs: leopard Panthera pardus diet in a human-dominated landscape in western Maharashtra, India

The ecology and predator–prey dynamics of large felids in the tropics have largely been studied in natural systems where wild ungulates constitute the majority of the prey base. However, human-dominated landscapes can be rich in potential prey for large carnivores because of the high density of domestic animals, especially in tropical countries where pastoralism is an important livelihood activity. We report the almost complete dependence of leopards Panthera pardus on domestic animals as prey in the crop lands of Ahmednagar district, Maharashtra, India. From analysis of 85 confirmed leopard scats, 87% of the leopards prey biomass consisted of domestic animals, with 39% consisting of domestic dogs Canis lupus familiaris alone. The only wild species that occurred in the leopards diet were rodents, small indian civet Viverricula indica , bonnet macaque Macaca radiata and other primates Semnopithecus spp., mongoose Herpestes spp., and birds. Interviews conducted in 77 households distributed randomly in the study area documented a high density of domestic animals: adult cattle Bos taurus , calves, goats Capra aegagrus , dogs and cats Felis catus occurred at densities of 169, 54, 174, 24 and 61 per km 2 , respectively. Ivlevs electivity index indicated that dogs and cats were over-represented in the leopards diet, given the higher densities of goats and cattle. The standing biomass of dogs and cats alone was sufficient to sustain the high density of carnivores at the study site. Our results show that the abundance of potential domestic prey biomass present in human-use areas supports a relatively high density of predators, although this interaction could result in conflict with humans.

Adaptable Neighbours: Movement Patterns of GPS-Collared Leopards in Human Dominated Landscapes in India

Understanding the nature of the interactions between humans and wildlife is of vital importance for conflict mitigation. We equipped five leopards with GPS-collars in Maharashtra (4) and Himachal Pradesh (1), India, to study movement patterns in human-dominated landscapes outside protected areas. An adult male and an adult female were both translocated 52 km, and exhibited extensive, and directional, post release movements (straight line movements: male  = 89 km in 37 days, female  = 45 km in 5 months), until they settled in home ranges of 42 km2 (male) and 65 km2 (female). The three other leopards, two adult females and a young male were released close to their capture sites and used small home ranges of 8 km2 (male), 11 km2 and 15 km2 (females). Movement patterns were markedly nocturnal, with hourly step lengths averaging 339±9.5 m (SE) during night and 60±4.1 m during day, and night locations were significantly closer to human settlements than day locations. However, more nocturnal movements were observed among those three living in the areas with high human population densities. These visited houses regularly at nighttime (20% of locations <25 m from houses), but rarely during day (<1%). One leopard living in a sparsely populated area avoided human settlements both day and night. The small home ranges of the leopards indicate that anthropogenic food resources may be plentiful although wild prey is absent. The study provides clear insights into the ability of leopards to live and move in landscapes that are extremely modified by human activity.

Trade-offs between maternal foraging and fawn predation risk in an income breeder

The choice of neonatal hiding place is critical for ungulates adopting hiding anti-predator strategies, but the consequences of different decisions have rarely been evaluated with respect to offspring survival. First, we investigated how landscape-scale choices made by roe deer fawns and their mothers affected predation risk by red foxes in a forest–farmland mosaic in southeastern Norway. After, we examined the effect of site-specific characteristics and behaviour (i.e. visibility, mother–fawn distance and abundance of the predator’s main prey item—small rodents) on predation risk. The study of habitat use, selection and habitat-specific mortality revealed that roe deer utilised the landscape matrix in a functional way, with different habitats used for feeding, providing maternal care and as refugia from predation. Mothers faced a trade-off between foraging and offspring survival. At the landscape-scale decisions were primarily determined by maternal energetic constraints and only secondarily by risk avoidance. Indeed, forage-rich habitats were strongly selected notwithstanding the exceptionally high densities of rodents which increased fawn predation. At fine spatial scales, a high visibility of the mother was the major factor determining predation risk; however, mothers adjusted their behaviour to the level of risk at the bed site to minimise predation. Fawns selected both landscape-scale refugia and concealed bed sites, but failure to segregate from the main prey of red foxes led to higher predation. This study provides evidence for the occurrence of spatial heterogeneity in predation risk and shows that energetically stressed individuals can tackle the foraging-safety trade-off by adopting scale-dependent anti-predator responses.

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

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