Christian Dussault

Effects of sampling effort based on GPS telemetry on home-range size estimations

Home-range size is widely used in wildlife biology to assess animal-habitat relationships. But home-range size estimation largely depends on the estimator and sample size used. Using 3 different estimators (minimum convex polygons, fixed-kernels, clusters), we used data collected on moose (Alces alces), fitted with Global Positioning System (GPS) collars (which allow automatic recording of large data sets with fewer field constraints than VHF telemetry) to investigate to what extent increasing the number of locations affects home-range size estimations. Our results indicated that 100 to 300 locations per animal annually and 30 to 100 locations seasonally were needed to reach an asymptote. High biases occurred below this asymptotic value. Minimum convex polygons consistently underestimated home-range sizes, but fixed-kernel and cluster estimators followed variable trends that often overestimated home-range sizes. Low sampling efforts also affected the number of animals needed to ensure an adequate statistical power of analysis to compare space use between 2 groups. Despite the higher cost, our findings indicate that in most instances, GPS telemetry is better suited than conventional radiotelemetry to estimate home-range sizes precisely and accurately. Moreover, this tracking technique is not limited by meteorological constraints and allows for the collection of similar sample sizes for all tracked animals, which is of major importance for further comparisons of space use among individuals.

Cumulative effects of forestry on habitat use by gray wolf (Canis lupus) in the boreal forest

Forest harvesting involves the creation of roads and cutblocks, both of which can influence animal habitat use. We evaluated the cumulative effects of forestry on habitat selection by six packs of gray wolf (Canis lupus) widely distributed in Quebec’s boreal forest. Resource selection functions were used to evaluate cumulative effects at two levels. First, we studied how the response of wolves to roads and cutblocks varied within their home range (HR level) as a function of the local abundance of these habitat features. Second, we assessed whether differences in the response to roads and cutblocks observed among packs (inter-HR level) could be explained by variations in their average abundance among individual home ranges. At the HR level, we found that cumulative effects shaped habitat selection of wolves, and the nature of the effects varied during the year. For example, we detected a decrease in the selection of roads following an increase in local road density during the rendez-vous and the nomadic periods, but not during the denning period. At the inter-HR level, we found a functional response to logging activity only during the denning period. Packs with home ranges characterized by a larger proportion of recent cutblocks selected these cutblocks more strongly. We conclude that cumulative effects of logging activities occur at multiple levels, and these effects can have profound effects on habitat use by wolves, thereby influencing spatial predator–prey dynamics. Wildlife conservation and management in boreal ecosystems should thus account for cumulative impacts of anthropogenic features on animal distribution.

Temporal and spatial distribution of moose-vehicle accidents in the Laurentides Wildlife Reserve, Quebec, Canada

Abstract Deer-vehicle accidents are an increasing problem in many regions of the world. To elaborate effective mitigation measures, it is necessary to determine environmental factors associated with the occurrence of such accidents. The Lauren-tides Wildlife Reserve in Quebec, Canada, is a prime example of an area having a long-lasting problem of moose-vehicle accidents (MVAs). We tested the effect of the spatial and temporal variables most likely to influence MVAs in this area based on accidents recorded over a 13-year period. Data collected included the date and time of each collision as well as the position of the accident relative to the closest 1-km road marker. We calculated 10 variables to assess moose habitat suitability, moose density, topography and road design for every 1-km road segment. There was a total of 754 MVAs during 1990–2002. The period with the highest number of accidents was the second half of June but accident frequency remained relatively high from mid-May to late August. The risk of accident per vehicle was at least 2–3 times higher at night (when traffic volume was lowest) than during any other time of the day. Also, MVAs were over 42% more frequent on Fridays, when road traffic levels were highest. The probability of an MVA increased when air temperature and atmospheric pressure were high. The MVA rate increased with moose density, in the presence of at least one brackish pool (by 80%) and when a valley traversed the road (by 120%). Implications of our results in the choice of appropriate mitigation measures are discussed. Future work should aim to describe habitat characteristics at the actual collision site.

Avoidance of roads and selection for recent cutovers by threatened caribou: fitness-rewarding or maladaptive behaviour?

The impact of anthropogenic disturbance on the fitness of prey should depend on the relative effect of human activities on different trophic levels. This verification remains rare, however, especially for large animals. We investigated the functional link between habitat selection of female caribou (Rangifer tarandus) and the survival of their calves, a fitness correlate. This top-down controlled population of the threatened forest-dwelling caribou inhabits a managed forest occupied by wolves (Canis lupus) and black bears (Ursus americanus). Sixty-one per cent of calves died from bear predation within two months following their birth. Variation in habitat selection tactics among mothers resulted in different mortality risks for their calves. When calves occupied areas with few deciduous trees, they were more likely to die from predation if the local road density was high. Although caribou are typically associated with pristine forests, females selected recent cutovers without negative impact on calf survival. This selection became detrimental, however, as regeneration took place in harvested stands owing to increased bear predation. We demonstrate that human disturbance has asymmetrical consequences on the trophic levels of a food web involving multiple large mammals, which resulted in habitat selection tactics with a greater short-term fitness payoff and, therefore, with higher evolutionary opportunity.

Balancing Number of Locations with Number of Individuals in Telemetry Studies

Abstract The study of habitat selection usually compares assessments of habitat use to habitat availability. To investigate habitat selection of large mammals today, researchers must choose between a few very expensive Global Positioning System (GPS) telemetry collars that can provide many locations and several inexpensive very high frequency telemetry collars that will provide few numbers of locations (unless substantial resources are spent in the field). We investigated the effects of number of locations and sampled animals on the outcome of habitat-selection analyses. We evaluated whether tracking frequency and sample size of individuals influenced our ability to detect habitat selection. We used data obtained from adult female moose fitted with GPS collars to generate data sets simulating various sampling frequencies and sample sizes of individuals. Tracking schedules conformed to those commonly used in ungulate telemetry studies (1 location every 14, 7, or 3 d and 1 or 3 locations per d) as did animal sample sizes (between 8 and 20 individuals). We determined habitat use and availability at the landscape and home-range scales during summer–autumn and winter. Precision of habitat use and availability estimates did not improve markedly with increasing tracking frequency. Only results obtained with the least-intensive tracking schedule (1 location every 14 d) differed from those obtained with the other schedules and only in 25% of the cases. Above this threshold in tracking frequency, number of sampled animals was clearly more important than number of locations in detection of habitat selection. Our results indicated that habitat-selection analyses were more sensitive to inter- than intra-individual variability. Depending on study objectives, it may be more profitable to prioritize number of sampled individuals rather than number of locations per individual. We suggest methods allowing researchers to assess inter-individual variability while studying habitat selection.

Plastic response of fearful prey to the spatiotemporal dynamics of predator distribution

Ecological theory predicts that the intensity of antipredator responses is dependent upon the spatiotemporal context of predation risk (the risk allocation hypothesis). However, most studies to date have been conducted over small spatial extents, and did not fully take into account gradual responses to predator proximity. We simultaneously collected spatially explicit data on predator and prey to investigate acute responses of a threatened forest ungulate, the boreal caribou (Rangifer tarandus), to the spatiotemporal dynamics of wolf (Canis lupus) distribution during spring. Movement analysis of GPS-collared individuals from both species revealed high plasticity in habitat-selection decisions of caribou. Female caribou avoided open areas and deciduous forests and moved relatively fast and toward foraging areas when wolves were closer than 2.5 km. Caribou also avoided food-rich areas only when wolves were within 1 km. Our results bridge the gap between long-term perceived risk and immediate flight responses by revealing dynamic antipredator tactics in response to predator proximity.

Multi‐trophic resource selection function enlightens the behavioural game between wolves and their prey

1. Habitat selection strategies translate into movement tactics, which reckon with the predator-prey spatial game. Strategic habitat selection analysis can therefore illuminate behavioural games. Cover types at potential encounter sites (i.e. intersections between movement paths of predator and prey) can be compared with cover types available (i) within the area of home-range-overlap (HRO) between predator and prey; and (ii) along the path (MP) of each species. Unlike the HRO scale, cover-type availability at MP scale differs between interacting species due to species-specific movement decisions. Scale differences in selection could therefore inform on divergences in fitness rewarding actions between predators and prey. 2. We used this framework to evaluate the spatial game between GPS-collared wolves (Canis lupus) versus caribou (Rangifer tarandus), and wolf versus moose (Alces alces). 3. Changes in cover-type availability between HRO and MP revealed differences in how each species fine-tuned its movements to habitat features. In contrast to caribou, wolves increased their encounter rate with regenerating cuts along their paths (MP) relative to the HRO level. As a consequence, wolves were less likely to cross caribou paths in areas with higher percentage of regenerating cuts than expected based on the availability along their paths, whereas caribou had a higher risk of intersecting wolf paths by crossing these areas, relative to random expectation along their paths. Unlike for caribou, availability of mixed and deciduous areas decreased from HRO to MP level for wolves and moose. Overall, wolves displayed stronger similarities in movement decisions with moose than with caribou, thereby revealing the focus of wolves on moose. 4. Our study reveals how differences in fine-scale movement tactics between species create asymmetric relative encounter probabilities between predators and prey, given their paths. Increase in relative risk of encounter for prey and decrease for predators associated with specific cover types emerging from HRO to MP scale analysis can disclose potential weaknesses in current movement tactics involved the predator-prey game, such as caribou use of cutovers in summer-autumn. In turn, these weaknesses can inform on subsequent changes in habitat selection tactics that might arise due to evolutionary forces.

Reducing Moose–Vehicle Collisions through Salt Pool Removal and Displacement: an Agent-Based Modeling Approach

Between 1990 and 2002, more than 200 moose-vehicle collisions occurred each year in Quebec, including about 50/yr in the Laurentides Wildlife Reserve. One cause is the presence of roadside salt pools that attract moose near roads in the spring and summer. Using the computer simulation technique of agent-based modeling, this study investigated whether salt pool removal and displacement, i.e., a compensatory salt pool set up 100 to 1500 m away from the road shoulder, would reduce the number of moose–vehicle collisions. Moose road crossings were used as a proxy measure. A GPS (global positioning system) telemetry data set consisting of approximately 200,000 locations of 47 moose over 2 yr in the Laurentides Wildlife Reserve was used as an empirical basis for the model. Twelve moose were selected from this data set and programmed in the model to forage and travel in the study area. Five parameters with an additional application of stochasticity were used to determine moose movement between forest polygons. These included food quality; cover quality, i.e., protection from predators and thermal stress; proximity to salt pools; proximity to water; and slope. There was a significant reduction in road crossings when either all or two thirds of the roadside salt pools were removed, with and/or without salt pool displacement. With 100% salt pool removal, the reduction was greater (49%) without compensatory salt pools than with them (18%). When two thirds of the salt pools were removed, the reduction was the same with and without compensatory salt pools (16%). Although moose-vehicle collisions are not a significant mortality factor for the moose population in the Laurentides Wildlife Reserve, in areas with higher road densities, hunting pressure, and/or predator densities it could mean the difference between a stable and a declining population, and salt pool removal could be part of a good mitigation plan to halt population declines. This model can be used, with improvements such as spatial memory of salt pool locations and the addition of a road avoidance behavior, to assess the effectiveness of mitigation measures intended to reduce moose–vehicle collisions.

Inference from habitat‐selection analysis depends on foraging strategies

1. Several methods have been developed to assess habitat selection, most of which are based on a comparison between habitat attributes in used vs. unused or random locations, such as the popular resource selection functions (RSFs). Spatial evaluation of residency time has been recently proposed as a promising avenue for studying habitat selection. Residency-time analyses assume a positive relationship between residency time within habitat patches and selection. We demonstrate that RSF and residency-time analyses provide different information about the process of habitat selection. Further, we show how the consideration of switching rate between habitat patches (interpatch movements) together with residency-time analysis can reveal habitat-selection strategies. 2. Spatially explicit, individual-based modelling was used to simulate foragers displaying one of six foraging strategies in a heterogeneous environment. The strategies combined one of three patch-departure rules (fixed-quitting-harvest-rate, fixed-time and fixed-amount strategy), together with one of two interpatch-movement rules (random or biased). Habitat selection of simulated foragers was then assessed using RSF, residency-time and interpatch-movement analyses. 3. Our simulations showed that RSFs and residency times are not always equivalent. When foragers move in a non-random manner and do not increase residency time in richer patches, residency-time analysis can provide misleading assessments of habitat selection. This is because the overall time spent in the various patch types not only depends on residency times, but also on interpatch-movement decisions. 4. We suggest that RSFs provide the outcome of the entire selection process, whereas residency-time and interpatch-movement analyses can be used in combination to reveal the mechanisms behind the selection process. 5. We showed that there is a risk in using residency-time analysis alone to infer habitat selection. Residency-time analyses, however, may enlighten the mechanisms of habitat selection by revealing central components of resource-use strategies. Given that management decisions are often based on resource-selection analyses, the evaluation of resource-use strategies can be key information for the development of efficient habitat-management strategies. Combining RSF, residency-time and interpatch-movement analyses is a simple and efficient way to gain a more comprehensive understanding of habitat selection.

Linking trade-offs in habitat selection with the occurrence of functional responses for moose living in two nearby study areas

A species may modify its relative habitat use with changing availability, generating functional responses in habitat selection. Functional responses in habitat selection are expected to occur when animals experience trade-offs influencing their habitat selection, but only a few studies to date have explicitly linked functional responses to the underlying trade-offs faced by the animals. We used data from 39 female moose fitted with GPS telemetry collars in two nearby study areas in Canada to investigate if moose (1) were faced with a food/cover trade-off in habitat selection, as typically acknowledged in the literature, and (2) showed a functional response in their use of food/cover-rich habitats. We also examined how habitat selection patterns varied seasonally, and between study areas. The occurrence of functional responses varied strongly between study areas, and could not always be related to a measurable food/cover trade-off. Functional responses were observed more often in the study area where the environmental conditions were more severe (colder temperatures, higher precipitations, and lower food availability). Selection coefficients were also less variable among individuals in that study area, suggesting that severe environmental conditions may constrain individuals to a few selection tactics and promote the development of functional responses. Moose reacted to the availability of different habitat types in different seasons, reflecting the changing trade-offs faced by the animals. We found considerable behavioral differences between individuals from two adjacent study areas, and therefore recommend caution when extrapolating habitat selection results. We advocate for the wider use of functional responses to identify critical habitats for a species from a management or conservation perspective.