A habitat-based framework for grizzly bear conservation in Alberta

Grizzly bear (Ursus arctos L.) populations in Alberta are threatened by habitat loss and high rates of human-caused mortality. Spatial depictions of fitness would greatly improve management and conservation action. We are currently challenged, however, in our ability to parameterize demographic rates necessary for describing fitness, especially across gradients of human disturbance and for land cover types. Alternative approaches are therefore needed. We describe here a method of estimating relative habitat states and conditions as surrogates of fitness using models of occupancy and mortality risk. By combining occurrence and risk models into a two-dimensional habitat framework, we identified indices of attractive sinks and safe harbour habitats, as well as five habitat states: non-critical habitats, secondary habitats (low-quality and secure), primary habitats (high-quality and secure), secondary sinks (low-quality, but high risk), and primary sinks (high-quality and high risk). Primary sink or high attractive sink situations were evident in the foothills where bears were using forest edges associated with forestry and oil and gas activities on Crown lands, while primary habitats or safe harbour sites were most common to protected alpine/sub-alpine sites. We suggest that habitat states and indices be used for setting baseline conditions for management and comparison of habitat conditions over time and identification of grizzly bear conservation reserves. A no net loss policy of critical habitats could be used to maintain existing habitat conditions for landscapes threatened by human development. Under such a policy, conversions of primary habitat would require restoration of equivalent amounts of primary sinks through decommissioning of roads.     

Can natural disturbance-based forestry rescue a declining population of grizzly bears?

Forest managers are increasingly considering historic patterns of natural forest disturbance as a model for forest harvesting and as a coarse-filter ecosystem management tool. We evaluated the long-term (100-year) persistence of a grizzly bear population in Alberta, Canada using forest simulations and habitat modelling. Even with harvesting the same volume of timber, natural disturbance-based forestry resulted in a larger human footprint than traditional two-pass forestry with road densities reaching 1.39 km/km² or more than three times baseline conditions and suggested maximum levels of security for grizzly bears. Because bears favour young forests and edges where food resources are plentiful, a future shift to young forests and more edge habitat resulted in a 20% projected increase in habitat quality and a 10% projected increase in potential carrying capacity. Human-caused mortality risk, however, offset any projected gains in habitat and carrying capacity resulting in the loss of all secure, unprotected territories, regardless of forest harvest method, within the first 20-30 years of simulation. We suggest that natural disturbance-based forestry is an ill-suited management tool for sustaining declining populations of grizzly bears. A management model that explicitly considers road access is more likely to improve grizzly bear population persistence than changing the size of clear-cuts. In fact, large clear cuts might be counter productive for bears since a diversity of habitats within each bear’s home range is more likely to buffer against future uncertainties.     

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.     

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.     

Dynamic wildlife habitat models: Seasonal foods and mortality risk predict occupancy-abundance and habitat selection in grizzly bears

Most current wildlife habitat models, such as resource selection functions, typically assume a static environment, extrapolate poorly in space and time, and often lack linkages to population processes. We submit that more mechanistic habitat models that directly consider bottom-up resources affecting growth and reproduction (i.e., food) and top-down limitations affecting survival are needed to effectively predict habitat quality, especially in the presence of rapid environmental change. Here we present a general model for estimating potential habitat quality (relating to growth and reproduction) and realised habitat quality (accounting for survival) using basic knowledge of the species’ seasonal diet, predicted locations of food resource patches and regional patterns in mortality risk. We illustrate our model for a threatened population of grizzly bears in west-central Alberta. Bi-monthly potential habitat quality successfully predicted habitat selection by radio-collared grizzly bears, while multi-seasonal realised habitat quality predicted patterns in occupancy-abundance as measured from unique bears at hair-snag sites. Bottom-up resources therefore predicted patterns of habitat selection, while top-down processes (survival) were necessary to scale-up to population measures. We suggest that more direct measures of resources and environments that affect growth, reproduction and survival, as well as match the temporal scale of animal behaviour, be considered when developing wildlife habitat models.     

Protecting more than the wetland: The importance of biased sex ratios and habitat segregation for conservation of the Hine’s emerald dragonfly, Somatochlora hineana Williamson

Within species habitat use may depend on age, season or sex of an individual. The distribution of males and females may vary both temporally and spatially due to differences in the costs of reproduction and the distribution of critical resources. Conservation of a species requires knowledge of the habitat use of both sexes in order to predict the population size and protect all habitats that a species requires. Adult dragonfly populations often have highly male-biased sex ratios at the breeding habitat. This bias has been attributed to females using alternative habitats to avoid male harassment, or to high female mortality. We monitored adult Hine’s emerald dragonfly (Somatochlora hineana Williamson) populations, in breeding and non-breeding habitats in Door County, Wisconsin and found significant differences in habitat use between males and females. Males primarily used wetland habitats, while females primarily used dry meadows and marginal breeding habitats, only coming into wetlands to lay-eggs or find mates. We assessed food resources in the different habitats and found that high quality insect prey (primarily adult Diptera) were more available in the wetland habitat, indicating that these areas were likely a more productive foraging area for adult dragonflies. The fact that females appear to avoid the wetland habitat is consistent with the hypothesis that male harassment alters female distribution patterns. Consideration of the patterns of habitat use by S. hineana indicates the need to develop a broader understanding of the importance of non-wetland areas in the conservation of wetland species.     

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.     

The effects of traffic on the raccoon dog (Nyctereutes procyonoides viverrinus) and other mammals in Japan

We investigated the extent of raccoon dog roadkills, and the factors influencing them, using data from the National Expressways in Japan for 1987-1998. A high rate of raccoon dog roadkills occurred in western Japan, whereas a relatively low rate was typical of the central region and of Hokkaido. Kills on the National Expressways increased as traffic and the length of the road network increased between 1987 and 1998. The relationship between mortality and traffic volume was scale-dependent. The number of roadkills was positively correlated with traffic volume at national level, but negatively correlated with the amount of traffic along sections of an individual major road between exits. There were spring and autumn peaks, and possibly early morning and evening peaks, of mortality. Mortality was higher for roads in a cutting, where the road was closer to water, and where the roadside habitat was broad-leaved woodland. Coniferous forest was negatively associated with roadkills. The total numbers of roadkills on four types of roads (National Expressways, National, Prefectural, and Municipal roads), was estimated to be between 110,000 and 370,000 in 1998. We suggest some preventative measures which might mitigate road casualties (while remaining attentive to human safety). Roadkill data, after controlling for traffic volume, can be useful as a population trend index. Patterns for racoon dogs are compared with those for other species.     

Efficacy of road removal for restoring wildlife habitat: Black bear in the Northern Rocky Mountains, USA

Forest roads reduce habitat quality for wildlife, in part by increasing susceptibility to hunting and poaching. Road removal is an increasingly common strategy for restoring habitat; however, little is known about responses of wildlife to road removal versus other methods of road closure. We assessed effects of different types of road closure (gated, barriered, and recontoured) on black bear (Ursus americanus) frequency and habitat on 18 open and closed road pairs in the western USA. Over 4 years, 44 bears were photographed during 3545 camera-trap days. Bear frequency was significantly higher (2.4 versus 0.6/100 days, respectively) and human frequency was significantly lower (2.4 versus 361.6/100 days, respectively) on closed than on open roads. Additionally, abundance of fall foods was higher (23.9% and 12.8%, respectively) and line-of-sight (a measure of habitat security) shorter (54.9 versus 69.4 m, respectively) on closed compared to open roads. Bears were detected on closed but not on open roads during daytime, suggesting avoidance of humans. Among-road-treatment differences included significantly higher frequency of bears on recontoured than on gated or barriered roads (4.6, 1.6, and 0.5/100 days, respectively), and significantly higher cover of fall bear foods on recontoured than on gated or barriered roads (39.3%, 12.1% and 16.4%, respectively). Frequency of bears was negatively correlated with frequency of humans and line-of-sight distance and positively correlated with abundance of fall foods and hiding cover. Results suggest that while all types of road closure benefit sensitive wildlife, removal by recontour may be the most effective strategy for restoring habitat.     

Origin and Transport of Aeolian Sediment from New Clearcuts into Boreal Lakes,Northwestern Ontario,Canada

Movement of aeolian (wind-blown) inorganic sediment was indexed in and around new riparian and upland clearcuts in northwestern Ontario, Canada with bucket-style terrestrial litterfall traps. Aeolian sediment originated primarily from clearcuts, roads, and skid trails, where soil disturbance was high and forest hitterfall was low. Relatively small amounts of aeolian sediment were recovered from riparian buffer strips and upland forest. Aeolian sediment was observed to reach lakes adjacent to the clearcuts and may be responsible for elevated levels of littoral sedimentation measured in one study lake. However, the amounts of sediment deposition observed are not likely to cause important changes in water quality or benthic habitat for algae, invertebrates, or fish in these lakes.     

Assessing the impacts of roads in peri-urban reserves: Road-based fatalities and road usage by wildlife in the Royal National Park, New South Wales, Australia

For protected reserves set aside for conservation, the impact of roads and traffic on wildlife can be severe, particularly for those in the peri-urban environment. Often reserves possess many sealed roads that have regular traffic from tourists and local residents. As managerial bodies struggle to control the wide variety of threats to the fauna within these reserves, the loss of life on roads only compounds the precarious nature of wildlife survival in these disturbed environments. As a first step to addressing this concern in Australia, this study quantifies the fatalities of wildlife killed on roads within the Royal National Park in New South Wales, and estimates those wildlife species using roadside habitat in order to identify species susceptible to collisions. Modelling of fatality data indicated that mammals were most likely to be killed where forage was abundant on the roadside verge and where there was plenty of protective cover, while birds were most likely to be killed when the height of roadside vegetation was low. A number of collision hotspots were identified along the surveyed road that should be the target of mitigation efforts. The average speed of vehicles travelling within the park peaked at night. This is of particular concern as activity by Australian mammals tends to be greatest at night. The findings indicate that roads in peri-urban reserves have the potential to alter the movement of animals and impact on their populations through loss of life.     

Performance indices to identify attributes of highway crossing structures facilitating movement of large mammals

Studies assessing the efficacy of wildlife crossing structures often lead to spurious results because of their failure to address masking effects of confounding variables. Confounding variables include variation in human activity, density of crossing structures along the highway corridor, and equality of species' perceived access to each crossing structure. We investigated these issues for wide-ranging large carnivores and their prey species in Banff National Park, Alberta, using data obtained from systematic, year-round monitoring of 13 newly constructed crossing structures for wildlife (underpasses and overpasses) for 34 months post-construction. We standardized the first confounding variable by selecting crossing structures remote from areas of human activity. The second confounding variable we standardized by developing probability models of crossing structure usage assuming habitat homogeneity. We standardized the third confounding variable by developing species-specific, performance indices of crossing structures (=observed through passage usage-expected through passage usage). We regressed the species performance indices against 13 crossing structure variables encompassing structural, landscape, and human activity. Our results suggest that in absence of high human activity structural attributes best explained the performance indices for both large predator and prey species, while landscape and human-related factors were of secondary importance. Crossing structures that were high, wide and short in length strongly influenced passage by grizzly bears Ursus arctos, wolves Canis lupus, elk Cervus elaphus, and deer Odocoileus sp. More constricted crossing structures were favoured by black bears Ursus americanus and cougars Puma concolor. Distance to cover was the most important crossing structure landscape attribute for cougars (negative correlation) and was a significant factor determining passage for grizzly bears, elk and deer (all positive correlations). Our findings underscore the importance of: (a) integrating temporal and spatial variability a priori when addressing the efficacy of crossing structures, and; (b) demonstrate that species respond differently to crossing structure features. In light of these results, we suggest that to maximize connectivity across roads for multiple large mammal species, road construction schemes should include a diversity of crossing structures of mixed size classes. Mitigation planning in a multiple-species ecosystem is likely to be a challenging endeavour and long-term research will aid in the decision-making process.     

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.     

Modelling the spatial distribution of human-caused grizzly bear mortalities in the Central Rockies ecosystem of Canada

We examined the spatial patterns of 297 human-caused grizzly bear mortalities from 1971 to 2002 within the Central Rockies ecosystem (CRE) of Canada to explore relationships between mortalities and variables reflecting human development, terrain, and vegetation. Using logistic regression, we modelled the distribution of grizzly bear mortalities based on local landscape attributes as well as examining variation among demographic status, seasons, and mortality type. Grizzly bear mortalities were concentrated in three main regions of the CRE: (1) Lake Louise; (2) Banff town site; and (3) Alberta Provincial lands near the Red Deer River. We found no evidence for environmental differences in mortality locations between sexes or seasons, while sub-adult male and legal harvest mortalities were more dispersed than other mortalities. Models describing the relative risk of mortality were positively associated with human access, water, and edge features, while negatively associated with terrain ruggedness and greenness indices. Model predictions fit well with independent data. Overall, relatively little of the landscape was secure from human-caused mortality for grizzly bears. This would be most directly remedied by controlling human access.     

Black bear resource selection in the northeast Cascades, Washington

We examined resource selection of black bears in the northeast Cascades of Washington at two spatial scales. Specifically, we compared habitats selected for within home ranges to those available in the study area, and habitats selected for versus those available within home ranges. Compositional analysis showed selection of similar habitats at each spatial scale, with some differences. In the dry climate of the eastern Cascades, black bears appeared to locate home ranges within habitats that would provide abundant food resources, such as riparian and deciduous forests, meadows and shrubfields. Once established in a home range, black bears selected for a mosaic of habitat types that provided security cover in proximity to food resources, such as riparian and deciduous forests, other forest types and meadows.     

A test of the hierarchical model of habitat selection using eastern massasauga rattlesnakes (Sistrurus c. catenatus)

Understanding the process animals follow to select habitat, rather than just documenting the habitat they use, will improve our ability to predict how the animals use habitat in other locations and how they will respond to changes in habitat. Animals are usually assumed to select habitats hierarchically, preferentially using specific macrohabitats at a landscape scale and specific microhabitats within the preferred macrohabitats. We used four years of telemetry data from 34 individuals to test this hierarchical model of habitat selection with eastern massasauga rattlesnakes (Sistrurus c. catenatus) in Ontario. Snakes were selective at the microhabitat scale, preferentially using locations with closer retreat sites and shrubs than random. Gravid females were most selective, using sites with more rock cover and less canopy closure than sites used by males and nongravid females. Snakes preferred forested habitats for hibernation and steadily increased their use of open, wetland, and edge habitats to a peak in mid-summer. Landscape-scale habitat preferences were generally mild and could be explained by the relative availability of suitable microhabitat within habitats, suggesting habitat selection was primarily driven by microhabitat preferences. The lack of selectivity at the landscape scale may be a consequence of fine-grained differences between habitats that allow massasauga rattlesnakes to find suitable microhabitats in all available macrohabitats. For species that select habitat primarily at the microhabitat scale (e.g., the rattlesnakes we studied), landscape-scale modeling of habitat use will only be effective to the extent habitats reflect the availability of suitable microhabitat within.     

Vigilance behaviour of polar bears (Ursus maritimus) in the context of wildlife-viewing activities at Churchill, Manitoba, Canada

Viewing of polar bears (Ursus maritimus) from tundra vehicles has been offered at Churchill, Manitoba since the early 1980s. This form of wildlife viewing has provided a unique and safe way for tourists to learn about polar bears. However, these activities have largely been carried out without examining possible effects on polar bear behaviour. We studied vigilance behaviour (a scanning of the immediate vicinity and beyond) of resting polar bears to evaluate impacts from tundra vehicle activity. Focal animal sampling was used to examine whether a difference in vigilance behaviour existed when vehicles were present. We recorded the numbers of head-ups, vigilance bout length, and between-bout intervals for polar bears. In general, the frequency of head-ups increased, and the between-bout intervals decreased for male bears, when vehicles were present. Female bears behaved opposite to males. The vigilance bout lengths did not differ significantly between vehicle presence and absence. Vigilance behaviour of male bears was not magnified with increasing numbers of vehicles; therefore the threshold is one vehicle. We suggest that manipulative studies be conducted to examine how distances between vehicles and bears, tundra vehicle activity in the immediate vicinity of a bear during viewing, and noise of tourists affect increased vigilance.     

Influence of the proximity and amount of human development and roads on the occurrence of the red imported fire ant in the lower Florida Keys

We examined the influence of both the proximity and extent of human developments and paved roads on the presence of the predatory, non-indigenous, red imported fire ant (Solenopsis invicta). This species was inadvertently introduced into the United States at the port of Mobile, Alabama, around 1930 and rapidly spread to many southeastern states, including Florida. More recently, S. invicta colonized the Florida Keys, an area with a high proportion of rare and endemic vertebrate and invertebrate species. We placed bait transects in transitional salt-marsh, pineland, and hardwood hammocks on 13 of the lower Florida Keys and compared habitat type, the shortest distance of the bait transect to a development or road, and area of development and roads 50, 70, 100, and 150 m around each bait transect for areas with and without red imported fire ants. Red imported fire ants were detected on 21 of the 80 transects and were equally abundant in all habitat types. While all of the development and road variables differed significantly between bait transects with and without red imported fire ants, transects that were closest to roads and that had the largest amount of development within a 150 m radii had the highest probability of presence of red imported fire ants. Recovery efforts for endangered species in areas invaded by red imported fire ants should include analyses of the cumulative impacts of roads and developments in areas near protected lands.     

Beyond the wetland border: Estimating the impact of roads for two species of water snakes

We used models integrating road maps, traffic volume, and snake movements to examine the potential for roads to contribute to mortality in two species of water snakes that differ in their vagility, use of terrestrial habitats, and conservation status. Road networks and traffic volumes typical of three regions in Indiana, USA, may account for mortality of 14-21% of the population per year in the more vagile, terrestrial, and imperiled copperbelly water snake (Nerodia erythrogaster neglecta) but only 3-5% mortality in the more sedentary, aquatic, and common northern water snake (Nerodia sipedon). The majority (>91%) of road crossings and associated mortality are predicted to occur during overland migrations to other wetlands, suggesting roads bisecting travel routes between wetlands may function as mortality sinks. Our models highlight the proportionately greater risk of mortality for the more vagile and imperiled species, N. e. neglecta, and suggest current wetland conservation strategies that focus on the wetland alone are unlikely to adequately protect wetland biodiversity from certain types of anthropogenic habitat modification. What is needed is a landscape approach to wetland conservation that considers not only the quality of wetlands and nearby terrestrial habitats, but also ensures that terrestrial corridors between wetlands remain permeable and offer safe passage for wildlife.