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.

Summary1. Caribou and reindeer Rangifer tarandus are declining across North America and Scandinavia in part from wolf Canis lupus-mediated apparent competition with more abundant ungulate prey species. While caribou generally persist in areas with low wolf density, wolf packs that overlap caribou ranges could trigger caribou declines. Moreover, anthropogenic linear features such as roads, trails and seismic lines are hypothesized to increase predation risk for caribou, yet few studies have examined the mechanistic effects of linear features or spatial overlap on wolf-caribou encounter rates and predation risk. 2. We used (a) time-to-event models of wolf-caribou encounters estimated from concurrent global positioning system (GPS) radio-collar data from wolves and caribou and (b) wolf resource selection models of travel locations, to determine the potential influence of wolf-caribou spatial overlap, linear features, elevation and season on encounter rates. Analyses were based on data from 35 adult female caribou and 37 male and female wolves from 11 wolf packs from Banff and Jasper National Parks, Canada, from 2002 until 2010. 3. Wolf-caribou encounter rates increased with high wolf-caribou overlap, proximity to linear features and lower elevations. Wolves strongly selected low elevations, especially during winter and spring. Selection for linear features as travel routes increased with elevation. 4. Caribou risk of encounter was highest during the summer and autumn when wolves spent the most time at high elevations. Most wolf-caused mortalities (n = 12) occurred during spring and summer. 5. Synthesis and applications. The presence of anthropogenic linear features and the amount of time wolves spend in caribou range could be equally as important as wolf density when prioritizing caribou recovery actions such as wolf or primary prey reductions or re-introductions. The use of GPS locations and time-to-event modelling offers a powerful tool for evaluating factors affecting predation risk of threatened and endangered species.

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.

Trends in population growth can be monitored with data for key vital rates without knowledge of abundance. Although adult female survival has the highest elasticity for ungulate population dynamics, the more variable recruitment rates are commonly monitored to track local variation in growth rates. Specifically, recruitment is often measured using late winter young:adult age ratios, though these age ratios are difficult to reliably interpret given the contribution of multiple vital rates to annual ratios. We show that the supplementation of age ratio data with concurrent radio-telemetry monitoring of adult female survival allows both retrospective estimation of empirical population growth rates and the decomposition of recruitment-specific vital rates. We demonstrate the estimation of recruitment and population growth rates for 1 woodland caribou population using these methods, including elasticity and life-stage simulation analysis of the relative contribution of adult female survival and recruitment rates to variation in population growth. We show, for this woodland caribou population, that adult female survival and recruitment rates were nearly equivalent drivers of population growth. We recommend the concurrent monitoring of adult female survival to reliably interpret age ratios when managing caribou and other ungulates. ß 2011 The Wildlife Society.

Highlights d Ungulates moved to track forage in landscapes with wavelike spring green-up d Patterns of green-up explained where migratory behavior occurred in many ecosystems d At the species level, migrants and residents received equivalent foraging benefits d Movement tactics represent behavioral adaptations to specific landscapes