Pragmatic management increases a flagship species, the Himalayan brown bears, in Pakistan’s Deosai National Park

Deosai Plateau in Northern Pakistan was designated a national park to protect the largest remnant population of brown bears in Pakistan. The natural resources of this high elevation (3500-5000 m) park make a significant contribution to the livelihood of local and nomad communities. The present legislation excludes people from a park, which increases conflicts between management and local people. However, a pragmatic approach was adopted to involve people in conservation in Deosai. Community participation, achieved by recognizing rights and introducing incentives, reduced resistance against the conservation efforts, reduced grazing pressure in bear habitat and helped reduce poaching. The size of the brown bear population was set as an indicator of park success, and was monitored annually from 1993 through 2006. We observed a 5% annual growth of the brown bear population, suggesting that the conservation program has been successful due to a successful cooperation between an NGO, people, and the park management.The increase of the bear population is significant, because we observed an extremely low reproductive rate, due to late age of first reproduction (8.25 years), a long reproductive interval (5.7 years), and a small litter size (1.33). The reproductive rate of the Deosai population is the lowest yet documented for brown bear populations. Poor habitat quality, low quality food, high seasonality, and extreme weather conditions in the Himalaya probably explain the poor reproductive performance. Considering such low reproduction and known exchange of individuals with neighboring populations, we believe that the observed growth was a sum of reproduction and immigration. Brown bears are declining throughout South Asia and often have low-productive rates. Therefore, conservation efforts for brown bears in this region must target reducing human-caused mortalities, particularly of adult females. Involvement of people can increase efficiency in conservation, in addition to reducing cost and conflicts.     

Estimating habitat suitability and potential population size for brown bears in the Eastern Alps

According to the Habitats Directive of the European Union, a favorable conservation status for the brown bear (Ursus arctos) should be targeted at the population level in large contiguous habitats such as the Alps, the largest mountain range in Europe. However, in most of the Alps brown bears are extinct and habitat suitability in these areas is often questionable. For this paper, radio-tracking data from four projects with 42 individual bears was compiled to assess habitat suitability. Discrete-choice models with random bear effects were fitted and compared to results obtained from compositional analysis and logistic regression. Sound definition of the available area in the discrete-choice model turned out to be essential. Brown bears showed a preference for forested and steep habitats and an avoidance of roads.Results from the three approaches were used to predict habitat suitability across the entire range of the Eastern Alps. Minimum potential population size was projected based on observed densities in Trentino and Central Austria, and ranged from 1228 to 1625 individuals, with 518–686 mature bears. This would satisfy a favorable conservation status. The developed methodology also has wide applicability to quantification of habitat suitability and potential population size in other cases where species are at risk.     

Genetic tracking of the brown bear in northern Pakistan and implications for conservation

Asian bears face major threats due to the impact of human activities as well as a critical lack of knowledge about their status, distribution and needs for survival. Once abundant in northern Pakistan, the Himalayan brown bear (Ursus arctos isabellinus) has been exterminated in most of its former distribution range. It presently occurs sparsely, in small populations, the Deosai National Park supporting the largest isolate. This decline might imply a reduction in genetic diversity, compromising the survival of the population. Using a combination of fecal DNA analysis and field data, our study aimed at assessing the size and genetic status of the Deosai population and give guidelines for its conservation and management. Using fecal genetic analysis, we estimated the population to be 40-50 bears, which compares well with the field census of 38 bears. The northern Pakistani brown bear population may have undergone an approximate 200-300-fold decrease during the last thousand years, probably due to glaciations and the influence of growing human population. However, in spite of the presence of a bottleneck genetic signature, the Deosai population has a moderate level of genetic diversity and is not at immediate risk of inbreeding depression. Gene flow might exist with adjacent populations. We recommend careful monitoring of this population in the future both with field observations and genetic analyses, including sampling of adjacent populations to assess incoming gene flow. The connectivity with adjacent populations in Pakistan and India will be of prime importance for the long-term survival of Deosai bears.     

Home-field advantage accelerates leaf litter decomposition in forests

Several leaf litter decay studies have indicated that decomposition occurs more rapidly when litter is placed beneath the plant species from which it had been derived than beneath a different plant species (i.e. home-field advantage, HFA), although support for this notion has not been universal. We provide the first quantification of HFA in relation to leaf litter decomposition using published litter mass loss data from forest ecosystems in North America, South America, and Europe. Our findings indicate that HFA is widespread in forest ecosystems; on average litter mass loss was 8% faster at home than away. We hypothesize that HFA results from specialization of the soil biotic community in decomposing litter derived from the plant above it. Climate and initial litter quality data can be used to explain about 70% of the variability in litter decomposition at a global scale, leaving about 30% unexplained. We suggest that HFA be recognized as a factor that explains some of this remaining variability.     

Density and population size estimates for North Cascade grizzly bears using DNA hair-sampling techniques

We used non-invasive DNA hair-sampling and catch per unit effort (CPUE: grizzly bears detected per 1000 trap nights) to estimate relative density and population size for a threatened grizzly bear population in the North Cascade Ecosystem of Washington and British Columbia. We used linear, logistic, and linear through the origin regression analyses to estimate the relationship between catch per unit effort and grizzly bear density for seven other grizzly populations. One grizzly bear was detected during 5304 trap nights (CPUE=0.19) over 3 years in the North Cascades. This CPUE was much lower than in the other seven populations, including two threatened grizzly populations in the Cabinet-Yaak and Selkirk Mountain Ecosystems. The logistic model (curvilinear relationship) best fit the data (R²=0.927), and yielded density and population size estimates of 0.15 bears/100 km² (90% CI=0.03-0.71) and six bears (90% CI=1-27), respectively. Natural recovery seems unlikely for the North Cascade grizzly bear population because the population has a high likelihood of extinction due to demographic and environmental stochastic effects associated with extremely small population numbers. We recommend population augmentation. DNA hair-sampling and catch per unit effort models can be a useful method to evaluate relative densities and numbers of animals in small, threatened grizzly bear populations when sample sizes are too small to yield traditional mark-recapture analysis.     

A habitat model for brown bear conservation and land use planning in the central Apennines

We investigated the brown bear habitat suitability in an 8000 km² study area encompassing Abruzzo, Latium, and Molise regions in central-southern Italy. Based on long-term field surveys and published records, we classified bear habitat as occupied or unoccupied in 92 out of 320 sample squares (5 × 5 km). For each sample square 36 habitat variables were measured from topographic maps and Corine land-cover III level digital maps. The influence of habitat features on bear presence was investigated by multivariate and one-way analyses of variance and by logistic regression analysis. The logistic model correctly classified 95.5% of sample squares of bear presence and 93.8% of those where bears were absent. Average altitude, deciduous woodlands and ecotone length, showed a positive relationship with bear presence, whereas vineyard-olive groves and shrublands were negatively correlated with bear presence. No specific land management guidelines or strategies exist for bear conservation in central Italy, based on knowledge of habitat-population relationships. The landscape scale habitat model we developed could be useful to predict bear occurrence, to identify critical areas for a brown bear conservation strategy, and to enhance the arrangement of the protected areas network for the conservation of this species.     

Perceived risk, displacement and refuging in brown bears: positive impacts of ecotourism?

Ecotourism is a rapidly growing industry with unknown impacts on viewed wildlife that may require novel management action. We examined the impact of viewing activities on the behaviour of brown bears (Ursus arctos) in coastal British Columbia.Domination of the best feeding sites and human avoidance by large male bears has consistently been reported. We, however, saw displacement in time rather than space - during the viewing day large males were less active than at other times, while females with cubs tended to be more active.In each year, females with cubs spent similarly high proportions of their time fishing when people were present. In years with large male activity, less time was spent fishing when people were absent. When freed from the potential threat of large male bears, females with cubs showed no measurable impact of controlled human activity.Human presence at a feeding site impacts the behaviour of brown bears, but not as expected. Temporal avoidance of human activity by large males was observed; indications that they departed upon satiation, before the arrival of morning tours, however, suggests that there was little energetic impact. By displacing large males, viewing activities created a temporal refuge, enhancing feeding opportunities for subordinate age/sex classes. With the strong positive relationships between mean female mass and litter size, this may in turn increase population productivity.     

Monitoring rare or elusive large mammals using effort-corrected voluntary observers

Populations of rare or elusive large mammals are difficult to monitor, because they usually are secretive, solitary, occur at low densities, and have large home ranges. The global trend of generally decreasing large carnivore populations necessitates new, feasible, reliable, and cost-effective monitoring methods. We evaluate an index method developed for monitoring populations of moose (Alces alces) based on voluntarily and systematically collected observations from hunters, corrected for effort, for use in monitoring populations of large carnivores in Sweden. For our evaluation, we used independent estimates of minimum brown bear (Ursus arctos) densities from DNA-based scat surveys and brown bear distribution from mandatory reports from successful bear hunters. We verified that the index correctly reflected bear distribution. We also found strong linear relationships between the indices and the independent density estimates for bears at the scale of local management units (about 1000-2000 km²) in all three regional study areas (adjusted R² = 0.88-0.60). Our results suggest that systematic, effort-corrected reports of observed animals can be an alternative and accurate monitoring method for the conservation and management of large mammals occurring over large areas when large numbers of willing volunteers are available (effort >30,000 h).     

An evaluation of field and non-invasive genetic methods to estimate brown bear (Ursus arctos) population size

Estimates of population size and density are essential for successful management and conservation of any species. Although there are a variety of methods available for estimating abundance and density of populations, most studies rely on only one estimator and very few studies have compared and critically evaluated the adequacy and the cost of these methods. We used the brown bear (Ursus arctos) in south-central Sweden to compare the performance of three different methods of estimating population size, including methods based on conventional field data as well as on non-invasive genetic data. The method based on observations of females with cubs underestimated the true population size, as the estimates were below the number of unique genotypes determined from faecal data inside the study area. The best traditional method was based on observations of bears from a helicopter. The genetic method using the closed population MARK estimator, as recommended in a previous study, seemed to perform the best. We conclude that approximately 223 (188-282) bears were present in our 7328 km² study area during 2001 and 2002 and suggest that this hunted brown bear population has been relatively stable for about ten years. The non-invasive genetic method was less expensive than the most reliable traditional field method (a CMR method based on observations of bears from a helicopter), and preferable from an ethical point of view. We recommend that future studies using non-invasive genetic methods based on collected faecal samples should aim at collecting 2.5-3 times the number of faecal samples as the “assumed” number of animals.     

Dynamics of public attitudes toward bears and the role of bear hunting in Croatia

Successful carnivore conservation depends on public attitudes and acceptance levels of carnivores, and these are likely to change as circumstances change. Attitude studies repeated in time that can demonstrate such change are rare. Our study surveyed Croatian rural inhabitants in 2002 and in 2008 and analyzed their responses to detect a change in attitudes toward brown bears (Ursus arctos) over time. Important developments occurring in Croatia at the time of our research included a more centralized and more clearly defined bear management strategy, and an increase in the bear population. We constructed models to explain respondent’s value orientations, their level of perceived threat and their acceptance capacity for bears. Findings show that while value orientations and the overall level of perceived threat did not change over time, bear acceptance capacity was reduced. This suggests that the increase in the bear population and perhaps the more centralized bear management reduced respondents’ willingness to accept a larger bear population. We conclude that continuous public involvement in bear management is essential in order to maintain a feeling of control over the bear among the local population. Furthermore we argue that hunting is an important form of public involvement in the region, serving to reinforce existence and bequest values of the bear and increase its public acceptance.     

Dynamics of a black bear population within a desert metapopulation

Understanding metapopulation dynamics in large carnivores with naturally fragmented populations is difficult because of the large temporal and spatial context of such dynamics. We coupled a long-term database of visitor sighting records with an intensive 3-year telemetry study to describe population dynamics of recolonization by black bears (Ursus americanus) of Big Bend National Park in Texas during 1988-2002. This population, which occurs within a metapopulation in western Texas and northern Mexico, increased from a single pair of known breeding-age animals in 1988 to 29 bears (including 6 females of breeding age) in March 2000 (λ = 1.25/year). A migration and dispersal event in August-December 2000 reduced the population to 2 adult females and as few as 5-7 individuals. One-way movement distances from the study area during this event averaged 76 km for females (n = 7) and 92 km for males (n = 4), and 3 animals conducted migrations of at least 154, 178, and 214 km, respectively. Our observations exemplify the importance of stochastic events on demographics of small populations and highlight the potential scale of bear movement among montane islands of southwestern North America. They also provide insight into the use of dispersal data in parameterizing metapopulation models for large carnivores.     

Impact of conservation interventions on the dynamics and persistence of a persecuted leopard (Panthera pardus) population

There is an extraordinary assortment of technical approaches to conserving carnivore populations, but the effectiveness of conservation activities is rarely evaluated. Accordingly, we initiated a study to assess the impact of several conservation interventions on the dynamics and persistence of a leopard (Panthera pardus) population in Phinda Private Game Reserve, South Africa. These included revisions of the statutory systems that regulate problem animal control and trophy hunting, and we instituted a program intended to reduce human–leopard conflict in the region. We compared demographic rates of radiocollared leopards from two sampling periods: a pre-treatment period prior to intervention, and a treatment period after intervention. The average annual mortality rate of the population decreased from 0.401 ± 0.070 to 0.134 ± 0.016, with fewer leopards killed by humans and in intraspecific clashes after intervention. The overall reproductive output of the population increased in the treatment period, although annual cub production was higher in the pre-treatment period. This was mainly due to larger litter sizes prior to intervention, which may have been a strategy used by female leopards to offset high levels of infanticide. Results from camera-trap surveys and a Leslie-matrix model indicated an increase in annual population growth rate (λ) of 14–16% after the implementation of conservation measures. Our findings highlight the importance of addressing both the numerical and functional components of population dynamics when managing large carnivores exposed to hunting or persecution.     

Low genetic variation for Dendrobaena octaedra from Greenland compared to populations from Europe and North America: Refuge or selection?

The genetic relationship of 345 specimens of the parthenogenetic lumbricid Dendrobaena octaedra from Greenland, Canada and Europe were analysed by means of isozymes. The results showed that populations from Greenland were markedly different from Canadian and European populations, suggesting that dispersal between Greenland and the continents has been much more restricted in the past than dispersal between North America and Europe. This observation supports to the notion that Greenland populations have persisted for a long period and perhaps have survived the last glacial period in ice-free refugia. A highly significant positive correlation was seen between diversity measured either as mean haploid diversity or clonal diversity and mean temperature of the annual coldest month. These results indicate that temperature might cause selection in colder climates or that sexual processes in D. dendrobaena could have been active recently on an evolutionary time scale.     

Public attitudes towards brown bears (Ursus arctos) in Slovenia

In northern Slovenia a radical change in brown bear (Ursus arctos) management - from a policy of bear suppression to a policy of bear protection - resulted in a sharp increase in sheep predation by bears. In the bear core area in southern Slovenia, on the other hand, bears have always been present, cause little damage and are an important game species. This zoned management provided a quasi “natural experiment” to compare attitudes and knowledge between two areas of different bear history, management and damage level. Using a questionnaire survey we sampled 924 locals and 177 hunters in the two areas. Contrary to our expectation, we documented a very positive attitude towards bears, in both study areas and for both target groups. Apparently, regional differences in the damage level per se and the status as a game species are not the driving force shaping attitudes towards bears in Slovenia. The key factor in predicting the attitude towards bears was the perception of how harmful the bears are. Knowledge and socio-demographic factors were only of minor importance. Even though attitude towards bears was positive, support of the present policy of bear expansion was low.     

Landscape evaluation of female black bear habitat effectiveness and capability in the North Cascades, Washington

We used logistic regression to derive scaled resource selection functions (RSFs) for female black bears at two study areas in the North Cascades Mountains. We tested the hypothesis that the influence of roads would result in potential habitat effectiveness (RSFs without the influence of roads) being greater than realized habitat effectiveness (RSFs with roads). Roads consistently had a negative influence on black bear RSFs across seasons and study areas. Roads reduced habitat effectiveness during all seasons at both study areas and changes in the potential habitat values ranged from 1.7% to 16.9%. The greatest reduction in habitat values occurred during the early-season on the west-side study area due to high open road densities. These results support the hypothesis that roads reduce habitat effectiveness for black bears. The influence of roads could be reduced through road closures to reduce open road densities and limit traffic volumes. We then used the scaled RSFs in a habitat-based population model to assess the influences of timber harvest and roads on potential black bear population sizes. On the west-side study area the potential black bear population size was most influenced by moderate use roads and timber harvest during the early-season (41% reduction). On the east-side study area, low use roads had the greatest effect on potential black bear population during the early-season (10% reduction). During the late-season, in both study areas, roads had less influence on the potential population sizes as bears were able to access habitats away from roads. The habitat-population model provided reasonable estimates of bear densities compared to other study areas with similar habitats and could be extrapolated to estimate potential black bear populations in other areas with similar habitats. This approach may provide a useful link between the landscape ecology and population biology of black bears, and could eventually be useful in the development of habitat-based population viability analyses.     

Determining the spatial scale for conservation purposes - an example with grizzly bears

This study suggests procedures for determining the spatial scale for conservation guidelines for animals, giving an illustration with an analysis of grizzly bear habitat selection. Bear densities were sampled by identifying hairs at bait stations in British Columbia. Habitat variables were measured using remote sensing. Spatial scale was changed by varying the window size over which the variables were averaged. First, the spatial pattern of bears was studied, measuring the patchiness in bear densities at a variety of spatial scales, by calculating the correlation in bear densities between adjacent windows. This was repeated for the habitat variables. Finally, the overall interaction between bears and habitats was analysed, measuring the strength of habitat selection at different spatial scales. There are three domains of scale: at 2-4 km, bears and habitats are patchy, at 5-10 km, bears select for habitats, and at 40+ km, habitats are patchy and bears select for habitats. At scales of 40+ km, bears selected for: (i) higher slopes, or (ii) higher slopes, and some combination of more avalanche chutes, fewer roads and trees, higher elevations, and less logged land. Within 15 km areas, bears selected for 6 km areas that are either at higher elevations, or at higher elevations and had fewer trees. The relationship of conservation guidelines at different spatial scales should be determined by measuring and comparing hierarchical to non-hierarchical selection. The scales that bears select for habitats roughly correspond to the scales used in present grizzly bear conservation plans in British Columbia.     

Human caused mortality in the endangered Scandinavian wolverine population

The wolverine is a relatively long lived mustelid species with low average annual reproduction which in theory makes them sensitive to changes in survival rates. In a multi-year study we monitored 211 radio-marked wolverines to estimate age-specific survival rates and mortality causes in an endangered population. Our data suggests that poaching forms a substantial part of wolverine population dynamics in northern Scandinavia causing up to 60% of adult mortality. Average annual adult survival rate was 0.91. Male and female survival rates did not differ. Twenty-five adult wolverines were confirmed to have died during the study. Fifty-two percent of confirmed adult mortality was human caused. The most important cause of adult mortality was poaching (9). Annual adult survival was lower (0.86) when 15 adult resident wolverines that were assumed to have died due to poaching were included in the analyses. Natural mortality among subadults and adults was less frequent in our study population compared to North American wolverine populations. Adult survival was significantly lower during the snow season (December-May) than during the snow-free season (June-November), while natural mortality was more evenly distributed between seasons. We explain this by the frequency of poaching which is higher during snow season. Our results underline the need to frame the underlying human dimension factors behind poaching to facilitate conservation and management of endangered populations.     

Island differences in population size structure and catch per unit effort and their conservation implications for Komodo dragons

Species inhabiting archipelagos are often characterised by high levels of interpopulation divergence (e.g. size related traits). This divergence may, in turn, influence their life-history. To facilitate better management and conservation of the Komodo dragon (Varanus komodoensis), an island endemic, we identified demographic differences between two island populations in Komodo National Park, Indonesia. Comparison of data collected from dragon populations inhabiting Rinca Island and the much smaller Gili Motang Island indicated that between 1994 and 2004, the Komodo dragon population on Gili Motang significantly decreased its: (1) mean body mass, (2) body condition and (3) relative abundance. These results suggest that the numerically small Gili Motang population was oscillating downwards; in contrast, the Rinca Island population had been relatively stable. More importantly these results emphasize the necessity for managers of this priority conservation species to understand further the inherent functional differences among dragon populations to develop island specific management units. Current management practices (e.g. monitoring) instigated by Komodo National Park management ignore small island dragon populations and thus run the risk of being unable to detect adverse effects for populations that are potentially most prone to decline.     

Demographic consequences of anthropogenic influences: Florida black bears in north-central Florida

Anthropogenic habitat fragmentation poses a serious threat to conservation of large carnivores, due to their extensive movements and potential conflicts with humans. We studied the population ecology of Florida black bears (Ursus americanus floridanus) for 6 years in two study areas in north-central Florida: Ocala National Forest (ONF), a contiguous forested habitat, and an adjacent residential community of Lynne, a fragmented habitat with substantial human activities. We estimated age-specific survival and fecundity rates of bears using data from radio-collared bears, and parameterized and analyzed stage-structured matrix population models for the two study sites and also for data pooled from both sites. Annual survival rates of adult females were lower in Lynne (0.776 ± 0.074) than in ONF (0.966 ± 0.023). While cub survival rates were higher in Lynne (0.507 ± 0.135) than in ONF (0.282 ± 0.109), the rates at both sites were substantially lower than those reported for other black bear populations. Age-specific fecundities did not vary between sites. The asymptotic population growth rate for ONF was greater than one, whereas that for Lynne was less than one. Our results suggest that anthropogenic influences (primarily road density and vehicular traffic, through their effect on adult survival) can substantially affect the population dynamics of Florida black bears and other large carnivores with large home ranges. We recommend efforts such as constructing highway underpasses, which could reduce road-related mortalities, to ensure long-term persistence of Florida black bears facing threats from rapidly increasing human influences.