Nicholas J. DeCesare

Resolving issues of imprecise and habitat-biased locations in ecological analyses using GPS telemetry data

Global positioning system (GPS) technologies collect unprecedented volumes of animal location data, providing ever greater insight into animal behaviour. Despite a certain degree of inherent imprecision and bias in GPS locations, little synthesis regarding the predominant causes of these errors, their implications for ecological analysis or solutions exists. Terrestrial deployments report 37 per cent or less non-random data loss and location precision 30 m or less on average, with canopy closure having the predominant effect, and animal behaviour interacting with local habitat conditions to affect errors in unpredictable ways. Home-range estimates appear generally robust to contemporary levels of location imprecision and bias, whereas movement paths and inferences of habitat selection may readily become misleading. There is a critical need for greater understanding of the additive or compounding effects of location imprecision, fix-rate bias, and, in the case of resource selection, map error on ecological insights. Technological advances will help, but at present analysts have a suite of ad hoc statistical corrections and modelling approaches available—tools that vary greatly in analytical complexity and utility. The success of these solutions depends critically on understanding the error-inducing mechanisms, and the biggest gap in our current understanding involves species-specific behavioural effects on GPS performance.

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.

Survival in the Rockies of an endangered hybrid swarm from diverged caribou (Rangifer tarandus) lineages

In North America, caribou (Rangifer tarandus) experienced diversification in separate refugia before the last glacial maximum. Geographical isolation produced the barren‐ground caribou (Rangifer tarandus groenlandicus) with its distinctive migratory habits, and the woodland caribou (Rangifer tarandus caribou), which has sedentary behaviour and is now in danger of extinction. Herein we report on the phylogenetics, population structure, and migratory habits of caribou in the Canadian Rockies, utilizing molecular and spatial data for 223 individuals. Mitochondrial DNA analyses show the occurrence of two highly diverged lineages; the Beringian–Eurasian and North American lineages, while microsatellite data reveal that present‐day Rockies’ caribou populations have resulted from interbreeding between these diverged lineages. An ice‐free corridor at the end of the last glaciation likely allowed, for the first time, for barren‐ground caribou to migrate from the North and overlap with woodland caribou expanding from the South. The lack of correlation between nuclear and mitochondrial data may indicate that different environmental forces, which might also include human‐caused habitat loss and fragmentation, are currently reshaping the population structure of this postglacial hybrid swarm. Furthermore, spatial ecological data show evidence of pronounced migratory behaviour within the study area, and suggest that the probability of being migratory may be higher in individual caribou carrying a Beringian–Eurasian haplotype which is mainly associated with the barren‐ground subspecies. Overall, our analyses reveal an intriguing example of postglacial mixing of diverged lineages. In a landscape that is changing due to climatic and human‐mediated factors, an understanding of these dynamics, both past and present, is essential for management and conservation of these populations.

Effect of forest canopy on GPS‐based movement data

Abstract The advancing role of Global Positioning System (GPS) technology in ecology has made studies of animal movement possible for larger and more vagile species. A simple field test revealed that lengths of GPS-based movement data were strongly biased (P<0.001) by effects of forest canopy. Global Positioning System error added an average of 27.5% additional length to tracks recorded under high canopy, while adding only 8.5% to open-canopy tracks, thus biasing comparisons of track length or tortuosity among habitat types. Other studies may incur different levels of bias depending on GPS sampling rates. Ninety-nine percent of track errors under high canopy were ≤7.98 m of the true path; this value can be used to set the scale-threshold at which movements are attributed to error and not biologically interpreted. This bias should be considered before interpreting GPS-based animal movement data.

Linking habitat selection and predation risk to spatial variation in survival.

1. A central assumption underlying the study of habitat selection is that selected habitats confer enhanced fitness. Unfortunately, this assumption is rarely tested, and in some systems, gradients of predation risk may more accurately characterize spatial variation in vital rates than gradients described by habitat selection studies. 2. Here, we separately measured spatial patterns of both resource selection and predation risk and tested their relationships with a key demographic trait, adult female survival, for a threatened ungulate, woodland caribou (Rangifer tarandus caribou Gmelin). We also evaluated whether exposure to gradients in both predation risk and resource selection value was manifested temporally through instantaneous or seasonal effects on survival outcomes. 3. We used Cox proportional hazards spatial survival modelling to assess the relative support for 5 selection- and risk-based definitions of habitat quality, as quantified by woodland caribou adult female survival. These hypotheses included scenarios in which selection ideally mirrored survival, risk entirely drove survival, non-ideal selection correlated with survival but with additive risk effects, an ecological trap with maladaptive selection and a non-spatial effect of annual variation in weather. 4. Indeed, we found positive relationships between the predicted values of a resource selection function (RSF) and survival, yet subsequently incorporating an additional negative effect of predation risk greatly improved models further. This revealed a positive, but non-ideal relationship between selection and survival. Gradients in these covariates were also shown to affect individual survival probability at multiple temporal scales. Exposure to increased predation risk had a relatively instantaneous effect on survival outcomes, whereas variation in habitat suitability predicted by an RSF had both instantaneous and longer-term seasonal effects on survival. 5. Predation risk was an additive source of hazard beyond that detected through selection alone, and woodland caribou selection thus was shown to be non-ideal. Furthermore, by combining spatial adult female survival models with herd-specific estimates of recruitment in matrix population models, we estimated a spatially explicit landscape of population growth predictions for this endangered species.

Movements, Connectivity, and Resource Selection of Rocky Mountain Bighorn Sheep

Abstract Species that exist in naturally fragmented subpopulations can maintain long-term viability through interpopulation connectivity and recolonization of suitable habitat. We used radiotelemetry to study movements of 3 herds of bighorn sheep (Ovis canadensis) that recently colonized previously unoccupied parts of western Montana. These herds also provided a unique opportunity to compare resource-selection patterns in newly colonized habitats, and we used logistic regression in a global information system framework to generate predictive models for females in each herd. We detected relatively long (19- to 33-km) extra–home range movements by males in all 3 herds, and connectivity with nearby bighorn and domestic sheep herds. An information-theoretic approach to model selection revealed greater differences in resource selection among herds than anticipated. Initial evaluation of resource-selection models by resubstituting data showed excellent predictive accuracy (P ≤ 0.002), but testing models across sites gave mixed results, and in many cases, poor fit (0.001 ≤ P ≤ 0.960). High vagility of males and variability in resource selection by females suggests increased potential for future recolonization and connectivity.

Preferred habitat and effective population size drive landscape genetic patterns in an endangered species

Landscape genetics provides a framework for pinpointing environmental features that determine the important exchange of migrants among populations. These studies usually test the significance of environmental variables on gene flow, yet ignore one fundamental driver of genetic variation in small populations, effective population size, Ne. We combined both approaches in evaluating genetic connectivity of a threatened ungulate, woodland caribou. We used least-cost paths to calculate matrices of resistance distance for landscape variables (preferred habitat, anthropogenic features and predation risk) and population-pairwise harmonic means of Ne, and correlated them with genetic distances, FST and Dc. Results showed that spatial configuration of preferred habitat and Ne were the two best predictors of genetic relationships. Additionally, controlling for the effect of Ne increased the strength of correlations of environmental variables with genetic distance, highlighting the significant underlying effect of Ne in modulating genetic drift and perceived spatial connectivity. We therefore have provided empirical support to emphasize preventing increased habitat loss and promoting population growth to ensure metapopulation viability.

Seasonal Resource Selection of Canada Lynx in Managed Forests of the Northern Rocky Mountains

Abstract We investigated seasonal patterns in resource selection of Canada lynx (Lynx canadensis) in the northern Rockies (western MT, USA) from 1998 to 2002 based on backtracking in winter (577 km; 10 M, 7 F) and radiotelemetry (630 locations; 16 M, 11 F) in summer. During winter, lynx preferentially foraged in mature, multilayer forests with Engelmann spruce (Picea engelmannii) and subalpine fir (Abies lasiocarpa) in the overstory and midstory. Forests used during winter were composed of larger diameter trees with higher horizontal cover, more abundant snowshoe hares (Lepus americanus), and deeper snow compared to random availability; multilayer, spruce–fir forests provided high horizontal cover with tree branching that touched the snow surface. During winter, lynx killed prey at sites with higher horizontal cover than that along foraging paths. Lynx were insensitive to snow depth or penetrability in determining where they killed prey. During summer, lynx broadened their resource use to select younger forests with high horizontal cover, abundant total shrubs, abundant small-diameter trees, and dense saplings, especially spruce–fir saplings. Based on multivariate logistic-regression models, resource selection occurred primarily at a fine spatial scale as was consistent with a sight-hunting predator in dense forests. However, univariate comparisons of patch-level metrics indicated that lynx selected homogenous spruce–fir patches, and avoided recent clear-cuts or other open patches. Given that lynx in Montana exhibit seasonal differences in resource selection, we encourage managers to maintain habitat mosaics. Because winter habitat may be most limiting for lynx, these mosaics should include abundant multistory, mature spruce–fir forests with high horizontal cover that are spatially well-distributed.

Hierarchical Den Selection of Canada Lynx in Western Montana

Abstract We studied den selection of Canada lynx (Lynx canadensis; hereafter lynx) at multiple ecological scales based on 57 dens from 19 females located in western Montana, USA, between 1999 and 2006. We considered 3 spatial scales in this analysis, including den site (11-m-radius circle surrounding dens), den area (100-m-radius circle), and den environ (1-km radius surrounding dens). Lynx denned in preexisting sheltered spaces created by downed logs (62%), root-wads from wind-thrown trees (19%), boulder fields (10%), slash piles (6%), and live trees (4%). Lynx preferentially selected den sites with northeasterly aspects that averaged 24°. Average distance between dens of 13 females monitored in consecutive years was 2,248 m, indicating low den site fidelity. Lynx exhibited habitat selection at all 3 spatial scales. Based on logistic regression, den sites differed from the surrounding den areas in having higher horizontal cover and log volume. Abundant woody debris from piled logs was the dominant habitat feature at den sites. Lynx generally denned in mature spruce–fir (Picea–Abies) forests with high horizontal cover and abundant coarse woody debris. Eighty percent of dens were in mature forest stands and 13% in mid-seral regenerating stands; young regenerating (5%) and thinned (either naturally sparse or mechanically thinned) stands with discontinuous canopies (2%) were seldom used. Female lynx selected den areas with greater spruce–fir tree basal area, higher horizontal cover, and larger-diameter trees compared to random locations within their home range. Lynx selected den environs in topographically concave or drainage-like areas, and farther from forest edges than random expectation. Maintaining mature and mid-seral regenerating spruce–fir forests with high horizontal cover and abundant woody debris would be most valuable for denning when located in drainages or in concave, drainage-like basins. Management actions that alter spruce–fir forests to a condition that is sparsely stocked (e.g., mechanically thinned) and with low canopy closure (<50%) would create forest conditions that are poorly suitable for lynx denning.

Separating spatial search and efficiency rates as components of predation risk

Predation risk is an important driver of ecosystems, and local spatial variation in risk can have population-level consequences by affecting multiple components of the predation process. I use resource selection and proportional hazard time-to-event modelling to assess the spatial drivers of two key components of risk—the search rate (i.e. aggregative response) and predation efficiency rate (i.e. functional response)—imposed by wolves (Canis lupus) in a multi-prey system. In my study area, both components of risk increased according to topographic variation, but anthropogenic features affected only the search rate. Predicted models of the cumulative hazard, or risk of a kill, underlying wolf search paths validated well with broad-scale variation in kill rates, suggesting that spatial hazard models provide a means of scaling up from local heterogeneity in predation risk to population-level dynamics in predator–prey systems. Additionally, I estimated an integrated model of relative spatial predation risk as the product of the search and efficiency rates, combining the distinct contributions of spatial heterogeneity to each component of risk.