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.     

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.     

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.     

Behavioral responses of brown bears mediate nutritional effects of experimentally introduced tourism

To develop a conceptual understanding of the interaction between tourists and bears, this study examined the responses of brown bears (Ursus arctos) to experimentally introduced tourism at Douglas River in south-central Alaska within the framework of current predator-prey theory. Factors eliciting displacement and the nutritional consequences of behavioral changes were measured. GPS collars were deployed to track temporal and spatial resource use and scan and focal observations were used to quantify foraging efficiency, vigilance behavior, and bear use of various food resources. Total food intake was quantified by combining data on food resource use based on GPS locations and foraging efficiency was measured using spotting scopes from long distances. Seasonal food availability and quality were monitored and utilized as co-variates in comparisons between pre-treatment and treatment years. Though the bear population exhibited significant behavioral differences between the presence and absence of humans, changes in behavior generally mediated effects on total food intake. Adult males at a salt marsh viewing area were the only sex/age class exhibiting reduced food intake resulting from a 15% decline in foraging time when viewers were present. Salmon intake did not differ between pre-treatment and treatment years despite a 10% decline in time spent fishing when viewers were present. The tightly controlled experimental bear-viewing introduced in this study allowed bears to evaluate predation risk associated with human activity and optimally respond to minimize costs due to changes in foraging activities.     

Environmental factors and spatial scale influence shorebirds’ responses to human disturbance

The extent of a shorebird’s response to a human disturbance depends on the associated energetic or predation risk costs. These costs are influenced by a suite of environmental variables, operating at several temporal and spatial scales. Here, I measured prey availability, distance to forest cover, cloud cover, and wind speed, in addition to human and shorebird densities to examine how human presence affected habitat choice, relative to these environmental variables, at Pachena Beach, British Columbia. In a standardised experiment, I also approached feeding flocks to determine if environmental factors influenced the time take for shorebirds to resume feeding. Binary logistic models suggested that people did not displace shorebirds. Instead, shorebirds were preferentially selecting areas further from forest cover that may have had lower predation risk. The time taken for shorebirds to resume feeding after a human disturbance was greater in the morning and in areas of low prey availability. This suggests that shorebirds respond more to a disturbance when the foraging cost is lower indicating that behavioural responses may not necessarily reflect the potential fitness costs of human disturbance.     

Reproduction and pre-dispersal survival of Iberian lynx in a subpopulation of the Doñana National Park

Little is known about the reproduction of the Iberian lynx (Lynx pardinus) even though it is the most endangered felid in the world. We studied during 9 years several reproductive parameters of the Iberian lynx in one of the subpopulations situated in Doñana National Park (south-western Spain), by means of radio-tracking, direct observations and photo-trapping. The potential breeding subpopulation was usually composed of 3 adult females, which bred 83% of the total possible 29 female-reproductive year. The minimum total number of cubs born during the study was 64 (7.7 ± 0.69 per year). There was no correlation between the number of cubs born or number of breeding females and population size of European rabbit (Oryctolagus cuniculus, the main prey of the lynx). All known births (n = 16) occurred in March except one in April and another in June. Mean litter size was 3.0 ± 0.16 (n = 16, range = 2-4). Sex-ratio (n = 59 cubs) was 1.03:1.00 (females:males). Nevertheless, the commonest picture was a female with 2 cubs older than 3 months. Altogether, at age of 3 months, 75% of cubs survived. Number of lynx alive at 10 months old and before dispersing was 69% and 57%, respectively. Sex did not affect survival for any age. Normally, it was not possible to know the causes of the death of cubs younger than 3 months. Survival at this age was not related with mother, mother age, or rabbit abundance. All females that bred were older than 3 years. The age of last reproduction was 9 years. For 3 females that were tracked during almost their complete reproductive life, the life time reproductive output was between 11 and 19 cubs. Iberian lynx reproductive parameters did not respond to wide changes in prey abundance during the study. Conservation plans considering the extraction of cubs with a low survival probability should be considered by managers, for instance, in translocation campaigns.     

Fish kills of Atlantic salmon (Salmo salar) and brown trout (Salmo trutta) in an acidified river of SW Norway

This paper documents population responses of Atlantic salmon and brown trout to fish kills in River Vikedal, an acidified river in SW Norway. The angling statistics show that the adult population of Atlantic salmon in the river has decreased to low numbers during recent years, whereas catches of migrant brown trout have increased in the same period. A total of 659 dead salmon and trout were either collected or observed during spring snowmelt in the years 1982–1985. Parr suffered the highest mortality, and most large specimens. Only a few dead kelts were registered. In the springs of most unfavourable water quality and most severe episods of fish kills (1983 and 1984), salmon parr mortality were significantly higher than that of brown trout. Episodic death of parr in the spring is thought to be an important cause of the reduction in the adult stock of Atlantic salmon in the river during recent years.     

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.     

Brown bear conservation and the ghost of persecution past

Large carnivores, such as brown bears, are focal species for conservation efforts. Historically, brown bears were persecuted in Europe for centuries before their gradual elimination from much of Western Europe. In contrast, large carnivore populations in North America were eradicated within two centuries in the east and within a few decades in the west. After a change towards conservation-oriented management in the 20th century, many bear populations are again increasing on both continents. Europe is seemingly less suited (i.e. higher human densities, greater habitat alteration and landscape fragmentation) than in North America, however bears seem to respond faster to conservation measures in Europe. We analyzed ecological and historical factors that may affect differences in reproductive allocation (mean litter size in relation to mean adult female body mass) and help explain why different brown bear populations react differently to conservation measures. The results indicated that mean litter size increased significantly with mean adult female body mass and a long persecution history. Our results suggest that high and long-term rates of nonselective harvesting can change life-history traits of large mammals, as has also been shown by modeling, but only has been documented for morphological traits. Incidentally, this “ghost of persecution past” may have helped some brown bear populations to be more productive and therefore to respond more positively to protective management policies than populations with short exploitation histories.     

Effects of Acidification on Salmonid Spawning Behavior

We studied the effects of acidification on female sexual behavior in brown trout (Salmo trutta) and compared the results with those in hime (land-locked sockeye) salmon (Oncorhynchus nerka) (Kitamura and Ikuta, 2000). The results were similar to those of sockeye salmon. Spawning brown trout were extremely sensitive to the acidity of ambient water, and nest-digging behavior was severely inhibited (p<0.05) by very slight acidification (pH below 6.4). However, there were some differences between the two species. Female trout and salmon showed almost no digging below pH 5.0 and 6.0 (Kitamura and Ikuta, 2000), respectively. When the ambient water was returned to nearly neutral (pH6.6) conditions, digging in hime salmon reappeared in 4 of the 6 fish tested (Kitamura and Ikuta, 2000), whereas digging in brown trout reappeared in all 6 fish tested. The above-mentioned differences in behavioral response between the two species appear to reflect the species difference in terms of vulnerability to acidification (Ikuta et al., 1992). Avoidance of slightly acidic water in selection of spawning site or cessation of spawning behavior in weakly acidic environments may be the most potent cause of the reduction of salmonid populations in the early stages of acidification.     

Sex differences in population genetics, home range size and habitat use of the parti-colored bat (Vespertilio murinus, Linnaeus 1758) in Switzerland and their consequences for conservation

When habitats are declining, niche segregation by demographic groups, such as the two sexes, can have a profound impact on the extinction risk of a species as a whole. Thus, differences in the requirements of demographic groups are of importance in conservation. We combined behavioural and genetic data to investigate whether the sexually segregated parti-colored bat (Vespertilio murinus) exhibits sex-specific niche partitioning. We use our data to evaluate implications for conservation of this potentially vulnerable species in Switzerland, the western boundary of its range. Using radio-telemetry, we found sex-specific differences in habitat use. Foraging females strongly relied on lakes while foraging males displayed more flexibility in their habitat use. Moreover, males covered significantly larger foraging areas than females. Sequencing 341 base pairs of the mitochondrial D-loop of 247 individuals revealed sex-specific differences in the genetic structure of colonies, but no such difference was observed for three nuclear micro-satellite markers. We found high mtDNA diversity in two Swiss male colonies and one German female colony, but low mtDNA diversity in two Swiss female colonies. Our genetic data suggest that considerable gene flow occurs via male dispersal and mating. At the same time immigration of females into the existing female colonies in Switzerland is rare compared to the immigration of new males into male colonies. Since we found the sexes in Vespertilio murinus to differ markedly in their ecology, population genetics, and behaviour, we conclude that sex-specific conservation plans are required to protect this species efficiently.     

The impact of sport-hunting on the population dynamics of an African lion population in a protected area

Between 1999 and 2004 we undertook an ecological study of African lions (Panthera leo) in Hwange National Park, western Zimbabwe to measure the impact of sport-hunting beyond the park on the lion population within the park, using radio-telemetry and direct observation. 34 of 62 tagged lions died during the study (of which 24 were shot by sport hunters: 13 adult males, 5 adult females, 6 sub-adult males). Sport hunters in the safari areas surrounding the park killed 72% of tagged adult males from the study area. Over 30% of all males shot were sub-adult (<4 years). Hunting off-take of male lions doubled during 2001-2003 compared to levels in the three preceding years, which caused a decline in numbers of adult males in the population (from an adult sex ratio of 1:3 to 1:6 in favour of adult females). Home ranges made vacant by removal of adult males were filled by immigration of males from the park core. Infanticide was observed when new males entered prides. The proportion of male cubs increased between 1999 and 2004, which may have occurred to compensate for high adult male mortality.     

Effects of supplementary feeding on the offspring sex ratio of kakapo: a dilemma for the conservation of a polygynous parrot

The kakapo (Strigops habroptilus) is a large, flightless, nocturnal parrot, endemic to New Zealand. It is critically endangered, with a world population of ca. 62 individuals and a male-biased adult sex ratio. The species has a polygynous “lek” mating system and adult males typically weigh 30-40% more than females. The kakapo is subject to intensive conservation efforts, including the provision of supplementary food to wild birds to encourage successful nesting. There is mounting evidence that, in polygynous species with large variance of male reproductive success, females in better condition may maximise their fitness by producing more offspring of the larger, more costly sex to be reared. We used data on the sex ratio of progeny of female kakapo that had or had not received supplementary food, to test the hypothesis that supplementary feeding might cause a male-biased offspring sex ratio. There was a significant excess of males in the clutches of females provided with supplementary food, suggesting that changes need to be made to the feeding regime to increase recruitment of females. This is an example of applying evolutionary theory to a practical conservation problem.     

Monitoring fish stocks in relation to acidification in Norwegian watersheds

The Norwegian Monitoring Programme for Long-Range Transported Air Pollutants started in 1980. The biological part of this programme includes besides invertebrate studies in streams, (i) fish community status in lakes by means of interviews, test-fishing in lakes by using standard gill-net series, recruitment studies of brown trout in inland streams, and juvenile stock assess and monitoring of fish kills in salmon rivers. Damaged fish stocks are recognized within a land area of 51,500 km² in southern Norway and 30 km² in northern Norway. At least 6,000 lake-dwelling fish stocks have either been lost or are at various stages of reduction. Brown trout (Salmo trutta) is the most widespread and abundant species of fish in Norwegian watersheds, and is also most severe affected by acidification. More recently, there are some indications of an increase in the abundance of brown trout in some areas. However, analysis of age structure in lakes, and fry densities in streams in such areas revealed large annual variations in recruitment rate, which indicates unstable water chemical conditions. Atlantic salmon (Salmo salar) is virtually extinct in 25 rivers in southernmost, southwestern and western Norway.     

The influence of microclimate on foraging and sheltering behaviours of terrestrial isopods: Implications for soil carbon dynamics under climate change

In south east England climate change is predicted to include decreases in periodicity of rainfall events during summer, resulting in more, longer periods of drought. In many temperate regions these changes will affect foraging strategies of soil animals, including arthropod macro-decomposers such as terrestrial isopods. We investigated activity budgets under drier (50%) and moisture (90%) relative humidity conditions, for three species: Porcellio scaber, Oniscus asellus and Armadillidium vulgare, representing three families, which have different morphological adaptations to the terrestrial environment. In the drier environment all three species spent more time sheltering, less time feeding and less time in other activities. Costs of foraging were estimated by monitoring loss of body mass for 7 h. This loss was twice as high under the lower humidity regime. On the basis of morphological traits, two members of the Oniscidae: O. asellus and Philoscia muscorum, differing in eco-morphological strategies, were predicted to be more susceptible to desiccation than members of the Porcellionidae or Amadillidiidae. Members of the Oniscidae lost mass most quickly and also suffered higher mortality in the drier atmosphere. A cost of spending more time sheltering was that consumption of high quality food by Porcellio scaber was significantly reduced in the drier environment. The trade-off between energy gain by feeding and increased probability of survivorship by sheltering, is illustrated using an isopleth model. Changes in the extent to which the soil fauna will stimulate microbial metabolism and hence soil carbon dioxide emissions, under future conditions of climate change are predicted.     

The impact of high speed, high volume traffic axes on brown bears in Slovenia

The Ljubljana-Razdrto highway and the parallel Ljubljana-Trieste railway cut through critical brown bear (Ursus arctos) habitat in south-central Slovenia. These high speed, high volume traffic axes are located close to the main dispersal corridor for bears from the Dinaric Mountain range into the Alps. We analyzed radiotracking data of 15 individual bears that lived within 10 km of the highway, compared transportation related and overall known bear mortality, and analyzed the spatial distribution of bear-vehicle accidents. The highway posed a home range boundary to resident bears, but was not an absolute barrier. Transportation-related mortality was high in the vicinity of the highway and railway, and averaged 31% of the total known local mortality from 1992 to 1999. At present the detrimental impact of transportation routes on the bear population in Slovenia is modest due to the high density of bears and the low density of highways—but new highways are planned or already under construction. Managers have to be aware that, due to bears large home ranges and long dispersal distances, a single highway affects bears from a huge area—emphasizing the importance of international cooperation and a landscape approach in highway planning.     

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.     

Evaluating estimators of the numbers of females with cubs-of-the-year in the yellowstone grizzly bear population

Current management of the grizzly bear (Ursus arctos) population in Yellowstone National Park and surrounding areas requires annual estimation of the number of adult female bears with cubs-of-the-year. We examined the performance of nine estimators of population size via simulation. Data were simulated using two methods for different combinations of population size, sample size, and coefficient of variation of individual sighting probabilities. We show that the coefficient of variation does not, by itself, adequately describe the effects of capture heterogeneity, because two different distributions of capture probabilities can have the same coefficient of variation. All estimators produced biased estimates of population size with bias decreasing as effort in creased. Based on the simulation results we recommend the Chao estimator for model M _h be used to estimate the number of the female bears with cubs of the year; however, the estimator of Chao and Shen may also be useful depending on the goals of the research.     

Location of roosts of the lesser long-eared bat Nyctophilus geoffroyi and Gould's wattled bat Chalinolobus gouldii in a fragmented landscape in south-eastern Australia

The location of roosts of two species of vespertilionid bats, the lesser long-eared bat (Nyctophilus geoffroyi) and Gould’s wattled bat (Chalinolobus gouldii), was investigated in a remnant vegetation-farmland mosaic and adjacent floodplain forest in south-eastern Australia. A total of 45 individuals of N. geoffroyi and 27 C. gouldii were fitted with radio transmitters, which resulted in the location of 139 and 89 roosts respectively. Selection of roosting habitat showed both inter- and intra-specific differences. At the landscape level, locations of roosts used by male and female N. geoffroyi differed markedly. Most males roosted in the farmland mosaic within 3 km (mean 1.9±2.9 km) of where they were captured while foraging in remnant vegetation. In contrast, roost sites of females were predominantly in the floodplain forest, 6-12 km from their capture site in farmland (mean for all females, 6.7±2.9 km). All maternity roosts were in the extensive floodplain forest, 4-10 km from foraging areas. Distances moved by this species were greater than previously recorded and further than predicted by flight morphology. Most C. gouldii roosted in the floodplain forest, 4-10 km from their capture site (6.9±1.6 km). Within the floodplain forest, both species roosted in areas that had higher densities of hollow-bearing trees than generally available; dead hollow-bearing trees for N. geoffroyi and large, live trees for C. gouldii. The scale of movements undertaken by these species means that they can obtain resources from multiple landscape elements. Both species used different habitats for roosting and foraging despite the energetic costs of commuting relatively large distances. Conservation of bat populations in rural environments needs to be considered at the landscape scale, with particular attention to identifying landscape elements that provide key resources.     

Baking reduces prostaglandin, resolvin, and hydroxy-fatty acid content of farm-raised Atlantic salmon (Salmo salar)

The consumption of seafood enriched in n-3 polyunsaturated fatty acids (PUFA) is associated with a decreased risk of cardiovascular disease. Several n-3 oxidation products from eicosapentaenoic acid (EPA; 20:5n-3) and docosahexaenoic acid (22:6n-3) have known protective effects in the vasculature. It is not known whether the consumption of cooked seafood enriched in n-3 PUFA causes appreciable consumption of lipid oxidation products. We tested the hypothesis that baking Atlantic salmon (Salmo salar) increases the level of n-3 and n-6 PUFA oxidation products over raw salmon. We measured the contents of several monohydroxy-fatty acids (MHFA), prostanoids, and resolvins. Our data demonstrate that baking did not change the overall total levels of MHFA. However, baking resulted in selective regioisomeric loss of hydroxy fatty acids from arachidonic acid (20:4n-6) and EPA, while significantly increasing hydroxyl-linoleic acid levels. The contents of prostanoids and resolvins were reduced several-fold with baking. The inclusion of a coating on the salmon prior to baking reduced the loss of some MHFA but had no effect on prostanoid losses incurred by baking. Baking did not decrease n-3 PUFA contents, indicating that baking of salmon is an acceptable means of preparation that does not alter the potential health benefits of high n-3 seafood consumption. The extent to which the levels of MHFA, prostanoids, and resolvins in the raw or baked fish have physiologic consequence for humans needs to be determined.