Corridor-based approach with spatial cross-validation reveals scale-dependent effects of geographic distance, human footprint and canopy cover on grizzly bear genetic connectivity.

Understanding how human infrastructure and other landscape attributes affect genetic differentiation in animals is an important step for identifying and maintaining dispersal corridors for these species. We built upon recent advances in the field of landscape genetics by using an individual-based and multiscale approach to predict landscape-level genetic connectivity for grizzly bears (Ursus arctos) across ~100,000 km2 in Canada's southern Rocky Mountains. We used a genetic dataset with 1156 unique individuals genotyped at nine microsatellite loci to identify landscape characteristics that influence grizzly bear gene flow at multiple spatial scales and map predicted genetic connectivity through a matrix of rugged terrain, large protected areas, highways and a growing human footprint. Our corridor-based modelling approach used a machine learning algorithm that objectively parameterized landscape resistance, incorporated spatial cross validation and variable selection and explicitly accounted for isolation by distance. This approach avoided overfitting, discarded variables that did not improve model performance across withheld test datasets and spatial predictive capacity compared to random cross-validation. We found that across all spatial scales, geographic distance explained more variation in genetic differentiation in grizzly bears than landscape variables. Human footprint inhibited connectivity across all spatial scales, while open canopies inhibited connectivity at the broadest spatial scale. Our results highlight the negative effect of human footprint on genetic connectivity, provide strong evidence for using spatial cross-validation in landscape genetics analyses and show that multiscale analyses provide additional information on how landscape variables affect genetic differentiation.

Changes in movement, habitat use, and response to human disturbance accompany parturition events in bighorn sheep (Ovis canadensis).

Parturition and the early neonatal period are critical life history stages in ungulates with considerable implications for population growth and persistence. Understanding the changes in behaviour induced by ungulate parturition is important for supporting effective population management, but reliably identifying birth sites and dates presents a challenge for managers. Rocky Mountain bighorn sheep (Ovis canadensis canadensis) are one such highly valued and ecologically important species in montane and subalpine ecosystems of Western North America. In the face of changing patterns of anthropogenic land use, wildlife managers increasingly require site-specific knowledge of the movement and habitat selection characteristics of periparturient sheep to better inform land use planning initiatives and ensure adequate protections for lambing habitat. We used movement data from GPS collared parturient (n = 13) and non-parturient (n = 8) bighorn sheep in Banff National Park, Canada to (1) identify lambing events based on changes in key movement metrics, and (2) investigate how resource selection and responses to human use change during the periparturient period. We fit a hidden Markov model (HMM) to a multivariate characterization of sheep movement (step length, daily home range area, residence time) to predict realistic lambing dates for the animals in our study system. Leave-one-out cross validation of our model resulted in a 93% success rate for parturient females. Our model, which we parameterized using data from known parturient females, also predicted lambing events in 25% of known non-parturient ewes in a test dataset. Using a latent selection difference function and resource selection functions, we tested for postpartum changes in habitat use, as well as seasonal differences in habitat selection. Immediately following lambing, ewes preferentially selected high-elevation sites on solar aspects that were more rugged, closer to escape terrain, and further from roads. Within-home range habitat selection was similar between individuals in different reproductive states, but parturient ewes had stronger selection for low snow depth, sites closer to barren ground, and sites further from trails. We propose that movement-based approaches such as HMMs are a valuable tool for identifying critical parturition habitat in species with complex movement patterns and may have particular utility in study areas without access to extensive field observations or vaginal implant transmitters. Furthermore, our results suggest that managers should minimize human disturbance in lambing areas to avoid interfering with maternal behaviour and ensure access to a broad range of suitable habitat in the periparturient period.

Protection status, human disturbance, snow cover and trapping drive density of a declining wolverine population in the Canadian Rocky Mountains.

Protected areas are important in species conservation, but high rates of human-caused mortality outside their borders and increasing popularity for recreation can negatively affect wildlife populations. We quantified wolverine (Gulo gulo) population trends from 2011 to 2020 in > 14,000 km2 protected and non-protected habitat in southwestern Canada. We conducted wolverine and multi-species surveys using non-invasive DNA and remote camera-based methods. We developed Bayesian integrated models combining spatial capture-recapture data of marked and unmarked individuals with occupancy data. Wolverine density and occupancy declined by 39%, with an annual population growth rate of 0.925. Density within protected areas was 3 times higher than outside and declined between 2011 (3.6 wolverines/1000 km2) and 2020 (2.1 wolverines/1000 km2). Wolverine density and detection probability increased with snow cover and decreased near development. Detection probability also decreased with human recreational activity. The annual harvest rate of ≥ 13% was above the maximum sustainable rate. We conclude that humans negatively affected the population through direct mortality, sub-lethal effects and habitat impacts. Our study exemplifies the need to monitor population trends for species at risk-within and between protected areas-as steep declines can occur unnoticed if key conservation concerns are not identified and addressed.

Towns and trails drive carnivore movement behaviour, resource selection, and connectivity.

BACKGROUND: Global increases in human activity threaten connectivity of animal habitat and populations. Protection and restoration of wildlife habitat and movement corridors require robust models to forecast the effects of human activity on movement behaviour, resource selection, and connectivity. Recent research suggests that animal resource selection and responses to human activity depend on their behavioural movement state, with increased tolerance for human activity in fast states of movement. Yet, few studies have incorporated state-dependent movement behaviour into analyses of Merriam connectivity, that is individual-based metrics of connectivity that incorporate landscape structure and movement behaviour. METHODS: We assessed the cumulative effects of anthropogenic development on multiple movement processes including movement behaviour, resource selection, and Merriam connectivity. We simulated movement paths using hidden Markov movement models and step selection functions to estimate habitat use and connectivity for three landscape scenarios: reference conditions with no anthropogenic development, current conditions, and future conditions with a simulated expansion of towns and recreational trails. Our analysis used 20 years of grizzly bear (Ursus arctos) and gray wolf (Canis lupus) movement data collected in and around Banff National Park, Canada. RESULTS: Carnivores increased their speed of travel near towns and areas of high trail and road density, presumably to avoid encounters with people. They exhibited stronger avoidance of anthropogenic development when foraging and resting compared to travelling and during the day compared to night. Wolves exhibited stronger avoidance of anthropogenic development than grizzly bears. Current development reduced the amount of high-quality habitat between two mountain towns by more than 35%. Habitat degradation constrained movement routes around towns and was most pronounced for foraging and resting behaviour. Current anthropogenic development reduced connectivity from reference conditions an average of 85%. Habitat quality and connectivity further declined under a future development scenario. CONCLUSIONS: Our results highlight the cumulative effects of anthropogenic development on carnivore movement behaviour, habitat use, and connectivity. Our strong behaviour-specific responses to human activity suggest that conservation initiatives should consider how proposed developments and restoration actions would affect where animals travel and how they use the landscape.

Disturbance type and species life history predict mammal responses to humans.

Human activity and land use change impact every landscape on Earth, driving declines in many animal species while benefiting others. Species ecological and life history traits may predict success in human-dominated landscapes such that only species with "winning" combinations of traits will persist in disturbed environments. However, this link between species traits and successful coexistence with humans remains obscured by the complexity of anthropogenic disturbances and variability among study systems. We compiled detection data for 24 mammal species from 61 populations across North America to quantify the effects of (1) the direct presence of people and (2) the human footprint (landscape modification) on mammal occurrence and activity levels. Thirty-three percent of mammal species exhibited a net negative response (i.e., reduced occurrence or activity) to increasing human presence and/or footprint across populations, whereas 58% of species were positively associated with increasing disturbance. However, apparent benefits of human presence and footprint tended to decrease or disappear at higher disturbance levels, indicative of thresholds in mammal species' capacity to tolerate disturbance or exploit human-dominated landscapes. Species ecological and life history traits were strong predictors of their responses to human footprint, with increasing footprint favoring smaller, less carnivorous, faster-reproducing species. The positive and negative effects of human presence were distributed more randomly with respect to species trait values, with apparent winners and losers across a range of body sizes and dietary guilds. Differential responses by some species to human presence and human footprint highlight the importance of considering these two forms of human disturbance separately when estimating anthropogenic impacts on wildlife. Our approach provides insights into the complex mechanisms through which human activities shape mammal communities globally, revealing the drivers of the loss of larger predators in human-modified landscapes.

Railway mortality for several mammal species increases with train speed, proximity to water, and track curvature.

Railways are a major source of direct mortality for many populations of large mammals, but they have been less studied or mitigated than roads. We evaluated temporal and spatial factors affecting mortality risk using 646 railway mortality incidents for 11 mammal species collected over 24 years throughout Banff and Yoho National Parks, Canada. We divided species into three guilds (bears, other carnivores, and ungulates), compared site attributes of topography, land cover, and train operation between mortality and paired random locations at four spatial scales, and described temporal patterns or mortality. Mortality risk increased across multiple guilds and spatial scales with maximum train speed and higher track curvature, both suggesting problems with train detection, and in areas with high proximity to and amount of water, both suggesting limitations to animal movement. Mortality risk was also correlated, but more varied among guilds and spatial scales, with shrub cover, topographic complexity, and proximity to sidings and roads. Seasonally, mortality rates were highest in winter for ungulates and other carnivores, and in late spring for bears, respectively. Our results suggest that effective mitigation could address train speed or detectability by wildlife, especially at sites with high track curvature that are near water or attractive habitat.

Temporal road closures improve habitat quality for wildlife.

Increasing levels of human activity threaten wildlife populations through direct mortality, habitat degradation, and habitat fragmentation. Area closures can improve habitat quality for wildlife, but may be difficult to achieve where tourism or other economic drivers are a priority. Temporal closures that limit human use during specific times of day have potential to increase habitat quality for wildlife, while continuing to provide opportunities for human use. However, the effectiveness of daily temporal closures has not been tested. We assessed how implementation of a temporal road closure affected wildlife movements in Banff National Park. Parks Canada closed a popular 17 km stretch of road between 2000 and 0800 hours to improve habitat quality for wildlife. We assessed the effectiveness of the closure on nine mammal species using three sets of data: remote cameras, road surveys, and grizzly bear (Ursus arctos) GPS data. In all three analyses, wildlife detection rates on the road doubled during the closure while remaining unchanged in reference areas. Our strong and consistent results suggest temporal closures are an important conservation tool that can increase habitat quality for wildlife while minimizing effects on people.

Design patterns for wildlife-related camera trap image analysis.

This paper describes and explains design patterns for software that supports how analysts can efficiently inspect and classify camera trap images for wildlife-related ecological attributes. Broadly speaking, a design pattern identifies a commonly occurring problem and a general reusable design approach to solve that problem. A developer can then use that design approach to create a specific software solution appropriate to the particular situation under consideration. In particular, design patterns for camera trap image analysis by wildlife biologists address solutions to commonly occurring problems they face while inspecting a large number of images and entering ecological data describing image attributes. We developed design patterns for image classification based on our understanding of biologists' needs that we acquired over 8 years during development and application of the freely available Timelapse image analysis system. For each design pattern presented, we describe the problem, a design approach that solves that problem, and a concrete example of how Timelapse addresses the design pattern. Our design patterns offer both general and specific solutions related to: maintaining data consistency, efficiencies in image inspection, methods for navigating between images, efficiencies in data entry including highly repetitious data entry, and sorting and filtering image into sequences, episodes, and subsets. These design patterns can inform the design of other camera trap systems and can help biologists assess how competing software products address their project-specific needs along with determining an efficient workflow.

Ecological correlates of the spatial co-occurrence of sympatric mammalian carnivores worldwide.

The composition of local mammalian carnivore communities has far-reaching effects on terrestrial ecosystems worldwide. To better understand how carnivore communities are structured, we analysed camera trap data for 108 087 trap days across 12 countries spanning five continents. We estimate local probabilities of co-occurrence among 768 species pairs from the order Carnivora and evaluate how shared ecological traits correlate with probabilities of co-occurrence. Within individual study areas, species pairs co-occurred more frequently than expected at random. Co-occurrence probabilities were greatest for species pairs that shared ecological traits including similar body size, temporal activity pattern and diet. However, co-occurrence decreased as compared to other species pairs when the pair included a large-bodied carnivore. Our results suggest that a combination of shared traits and top-down regulation by large carnivores shape local carnivore communities globally.

Sampling scales define occupancy and underlying occupancy-abundance relationships in animals.

Occupancy-abundance (OA) relationships are a foundational ecological phenomenon and field of study, and occupancy models are increasingly used to track population trends and understand ecological interactions. However, these two fields of ecological inquiry remain largely isolated, despite growing appreciation of the importance of integration. For example, using occupancy models to infer trends in abundance is predicated on positive OA relationships. Many occupancy studies collect data that violate geographical closure assumptions due to the choice of sampling scales and application to mobile organisms, which may change how occupancy and abundance are related. Little research, however, has explored how different occupancy sampling designs affect OA relationships. We develop a conceptual framework for understanding how sampling scales affect the definition of occupancy for mobile organisms, which drives OA relationships. We explore how spatial and temporal sampling scales, and the choice of sampling unit (areal vs. point sampling), affect OA relationships. We develop predictions using simulations, and test them using empirical occupancy data from remote cameras on 11 medium-large mammals. Surprisingly, our simulations demonstrate that when using point sampling, OA relationships are unaffected by spatial sampling grain (i.e., cell size). In contrast, when using areal sampling (e.g., species atlas data), OA relationships are affected by spatial grain. Furthermore, OA relationships are also affected by temporal sampling scales, where the curvature of the OA relationship increases with temporal sampling duration. Our empirical results support these predictions, showing that at any given abundance, the spatial grain of point sampling does not affect occupancy estimates, but longer surveys do increase occupancy estimates. For rare species (low occupancy), estimates of occupancy will quickly increase with longer surveys, even while abundance remains constant. Our results also clearly demonstrate that occupancy for mobile species without geographical closure is not true occupancy. The independence of occupancy estimates from spatial sampling grain depends on the sampling unit. Point-sampling surveys can, however, provide unbiased estimates of occupancy for multiple species simultaneously, irrespective of home-range size. The use of occupancy for trend monitoring needs to explicitly articulate how the chosen sampling scales define occupancy and affect the occupancy-abundance relationship.

Seasonal and individual variation in the use of rail-associated food attractants by grizzly bears (Ursus arctos) in a national park.

Similar to vehicles on roadways, trains frequently kill wildlife via collisions along railways. Despite the prevalence of this mortality worldwide, little is known about the relative importance of wildlife attractants associated with railways, including spilled agricultural products, enhanced vegetation, invertebrates, and carcasses of rail-killed ungulates. We assessed the relative importance of several railway attractants to a provincially-threatened population of grizzly bears (Ursus arctos) in Banff and Yoho National Parks, Canada, for which rail-caused mortality has increased in recent decades without known cause. We examined the relationship between the use of the railway and diet by fitting 21 grizzly bears with GPS collars in 2011-2013 and measuring the stable isotope values (δ15N, δ34S) derived from their hair. We also examined the importance of rail-associated foods to grizzly bears by analyzing 230 grizzly bear scats collected from May through October in 2012-2014, some of which could be attributed to GPS-collared bears. Among the 21 collared bears, 17 used the rail rarely (<9% of the days they were monitored), and only four bears (which included the three smallest bears and the largest bear in our sample) used the rail frequently (>20% of their monitored days). We found no significant relationships between δ15N and δ34S values measured from the hair of grizzlies and their frequency of rail use. Instead, δ15N increased with body mass, especially for male bears, suggesting large males consumed more animal protein during hair growth. All four bears that used the railway frequently produced scats containing grain. Almost half the scats (43%) collected within 150 m of the railway contained grain compared to only 7% of scats found >150 m from the railway. Scats deposited near the rail were also more likely to contain grain in the fall (85% of scats) compared to summer (14%) and spring (17%), and those containing grain were more diverse in their contents (6.8 ± 2.2 species vs. 4.9 ± 1.6, P < 0.001). Lastly, scats collected near the rail were more likely to contain ungulate hair and ant remains, especially in the summer. Our results support local management knowledge that some bears in the region use the railway to forage and supplement their diets with spilled grain, but that individual use of the railway and associated foods were highly variable. We suggest that managers continue to reduce the risk of bears being killed by trains by reactively removing grain and ungulate carcasses from the railway, reducing the amount of grain spilled by trains, and target mitigation to the specific individuals and locations that attract recurrent rail-based foraging.

Behavioural flexibility in migratory behaviour in a long-lived large herbivore.

Migratory animals are predicted to enhance lifetime fitness by obtaining higher quality forage and/or reducing predation risk compared to non-migratory conspecifics. Despite evidence for behavioural flexibility in other taxa, previous research on large mammals has often assumed that migratory behaviour is a fixed behavioural trait. Migratory behaviour may be plastic for many species, although few studies have tested for individual-level flexibility using long-term monitoring of marked individuals, especially in large mammals such as ungulates. We tested variability in individual migratory behaviour using a 10-year telemetry data set of 223 adult female elk (Cervus elaphus) in the partially migratory Ya Ha Tinda population in Alberta, Canada. We used net squared displacement (NSD) to classify migratory strategy for each individual elk-year. Individuals switched between migrant and resident strategies at a mean rate of 15% per year, and migrants were more likely to switch than residents. We then tested how extrinsic (climate, elk/wolf abundance) and intrinsic (age) factors affected the probability of migrating, and, secondly, the decision to switch between migratory strategies. Over 630 individual elk-years, the probability of an individual elk migrating increased following a severe winter, in years of higher wolf abundance, and with increasing age. At an individual elk level, we observed 148 switching events of 430 possible transitions in elk monitored at least 2 years. We found switching was density-dependent, where migrants switched to a resident strategy at low elk abundance, but residents switched more to a migrant strategy at high elk abundance. Precipitation during the previous summer had a weak carryover effect, with migrants switching slightly more following wetter summers, whereas residents showed the opposite pattern. Older migrant elk rarely switched, whereas resident elk switched more frequently to migrate at older ages. Our results show migratory behaviour in ungulates is an individually variable trait that can respond to intrinsic, environmental and density-dependent forces. Different strategies had opposing responses to density-dependent and intrinsic drivers, providing a stabilizing mechanism for the maintenance of partial migration and demographic fitness in this population.

A Comparison of Grizzly Bear Demographic Parameters Estimated from Non-Spatial and Spatial Open Population Capture-Recapture Models.

Capture-recapture studies are frequently used to monitor the status and trends of wildlife populations. Detection histories from individual animals are used to estimate probability of detection and abundance or density. The accuracy of abundance and density estimates depends on the ability to model factors affecting detection probability. Non-spatial capture-recapture models have recently evolved into spatial capture-recapture models that directly include the effect of distances between an animal's home range centre and trap locations on detection probability. Most studies comparing non-spatial and spatial capture-recapture biases focussed on single year models and no studies have compared the accuracy of demographic parameter estimates from open population models. We applied open population non-spatial and spatial capture-recapture models to three years of grizzly bear DNA-based data from Banff National Park and simulated data sets. The two models produced similar estimates of grizzly bear apparent survival, per capita recruitment, and population growth rates but the spatial capture-recapture models had better fit. Simulations showed that spatial capture-recapture models produced more accurate parameter estimates with better credible interval coverage than non-spatial capture-recapture models. Non-spatial capture-recapture models produced negatively biased estimates of apparent survival and positively biased estimates of per capita recruitment. The spatial capture-recapture grizzly bear population growth rates and 95% highest posterior density averaged across the three years were 0.925 (0.786-1.071) for females, 0.844 (0.703-0.975) for males, and 0.882 (0.779-0.981) for females and males combined. The non-spatial capture-recapture population growth rates were 0.894 (0.758-1.024) for females, 0.825 (0.700-0.948) for males, and 0.863 (0.771-0.957) for both sexes. The combination of low densities, low reproductive rates, and predominantly negative population growth rates suggest that Banff National Park's population of grizzly bears requires continued conservation-oriented management actions.

Stable isotopes reveal rail-associated behavior in a threatened carnivore.

Human-wildlife conflict is a leading cause of adult mortality for large carnivores worldwide. Train collision is the primary cause of mortality for threatened grizzly bears (Ursus arctos) in Banff National Park. We investigated the use of stable isotope analysis as a tool for identifying bears that use the railway in Banff. Rail-associated bears had higher δ(15)N and δ(34)S values than bears sampled away from the rail, but similar δ(13)C values. Because elevated δ(15)N values are indicative of higher animal protein consumption, rail-associated bears likely preyed on ungulates that foraged along the rail or scavenged on train-killed animals. The higher δ(34)S values in bear hair could have resulted from bears consuming sulfur pellets spilled on the rail or through the uptake of sulfur in the plants bears or animals consumed. Similar δ(13)C values suggest that the two types of bears had generally similar plant-based diets. Results from this study suggest that stable isotopes analysis could be used as a non-invasive, affordable, and efficient technique to identify and monitor bears that forage on the railway in Banff and potentially other transportation corridors worldwide.

Transcending scale dependence in identifying habitat with resource selection functions.

Multi-scale resource selection modeling is used to identify factors that limit species distributions across scales of space and time. This multi-scale nature of habitat suitability complicates the translation of inferences to single, spatial depictions of habitat required for conservation of species. We estimated resource selection functions (RSFs) across three scales for a threatened ungulate, woodland caribou (Rangifer tarandus caribou), with two objectives: (1) to infer the relative effects of two forms of anthropogenic disturbance (forestry and linear features) on woodland caribou distributions at multiple scales and (2) to estimate scale-integrated resource selection functions (SRSFs) that synthesize results across scales for management-oriented habitat suitability mapping. We found a previously undocumented scale-specific switch in woodland caribou response to two forms of anthropogenic disturbance. Caribou avoided forestry cut-blocks at broad scales according to first- and second-order RSFs and avoided linear features at fine scales according to third-order RSFs, corroborating predictions developed according to predator-mediated effects of each disturbance type. Additionally, a single SRSF validated as well as each of three single-scale RSFs when estimating habitat suitability across three different spatial scales of prediction. We demonstrate that a single SRSF can be applied to predict relative habitat suitability at both local and landscape scales in support of critical habitat identification and species recovery.

The role of translocation in recovery of woodland caribou populations.

Maintenance of viable populations of many endangered species will require conservation action in perpetuity. Efforts to conserve these species are more likely to be successful if their reliance on conservation actions is assessed at the population level. Woodland caribou (Rangifer tarandus caribou) were extirpated recently from Banff National Park, Canada, and translocations of caribou to Banff and neighboring Jasper National Park are being considered. We used population viability analysis to assess the relative need for and benefits from translocation of individuals among caribou populations. We measured stochastic growth rates and the probability of quasi extinction of four populations of woodland caribou with and without translocation. We used two vital rates in our analysis: mean adult female survival and mean number of calves per breeding-age female as estimates of mean fecundity. We isolated process variance for each vital rate. Our results suggested the Tonquin caribou population in Jasper is likely to remain viable without translocation, but that translocation is probably insufficient to prevent eventual extirpation of the two other populations in Jasper. Simulated reintroductions of caribou into Banff resulted in a 53-98% probability of >8 females remaining after 20 years, which suggests translocation may be an effective recovery tool for some caribou populations.