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.

Territorial behavior is expected to buffer populations against short-term environmental perturbations, but we have found that group living in African lions causes a complex response to long-term ecological change. Despite numerous gradual changes in prey availability and vegetative cover, regional populations of Serengeti lions remained stable for 10- to 20-year periods and only shifted to new equilibria in sudden leaps. Although gradually improving environmental conditions provided sufficient resources to permit the subdivision of preexisting territories, regional lion populations did not expand until short-term conditions supplied enough prey to generate large cohorts of surviving young. The results of a simulation model show that the observed pattern of "saltatory equilibria" results from the lions' grouping behavior.

Many animal species are unevenly distributed across the landscape, in spatial patterns that continually shift over time. Such a shifting mosaic is thought to have profound implications for the persistence and stability of ecosystems. Management and conservation of natural systems would be enhanced if we could accurately predict movement. Such prediction has not yet been possible. Here we use an extensive set of field data on food abundance and quality, combined with experimentally derived measures of nutritional value, to predict the spatial distribution of Thomson's gazelles (Gazella thomsoni thomsoni Gü nter) on the Serengeti Plains of East Africa. Twelve plausible models, based on alternate foraging objectives or movement rules, were assessed against field data on food and grazer abundance gathered at biweekly intervals (every two weeks) over the course of the wet seasons in two different years. Nomadic movements of gazelles closely tracked changes in the spatial distribution of short grass swards. Gazelles left short grass patches when local daily energy intake dropped below the expected intake averaged across the landscape. Subsequent redistribution of gazelles among neighboring patches was proportional to daily rates of energy intake in each patch. Thus, nomadic movements by Thomson's gazelles were predictable on the basis of local energy gain. This suggests that adaptive behavioral models can provide useful predictive tools for understanding the dynamics of complex natural systems.

An axiomatic feature of food consumption by animals is that intake rate and prey abundance are positively related. While this has been demonstrated rigorously for large herbivores, it is apparent from patch selection trials that grazers paradoxically tend to prefer short, sparse swards to tall, dense swards. Indeed, migratory herbivores often shift from areas of high to low sward biomass during the growing season. As nutritional quality is an inverse function of grass abundance, herbivores appear to sacri¢ce short-term intake for nutritional gains obtainable by eating sparse forage of higher quality. Explicit models of this trade-o¡ suggest that individual ruminants maximize daily rates of energy gain by choosing immature swards of intermediate biomass. As body mass is related positively to both ruminant cropping rates and digestibility, there should be an allometric link between grass abundance and energy maximization, providing a tool for predicting patterns of herbivore habitat selection. We used previously published studies to develop a synthetic model of trade-o¡s between forage abundance and quality, predicting that optimal sward biomass should scale allometrically with body size. The model predicts size-related variation in habitat selection observed in a guild of grazing ungulates in the Serengeti ecosystem.

Species persistence can be threatened by substantial temporal variation in food resources over time. On the other hand, spatial heterogeneity in resources at the landscape scale might allow mobile consumers to compensate for temporal variability in resource availability at the local scale. We evaluated this hypothesis, using an extensive data set on foraging, grass growth, and movement by Thomson's gazelles living on the Serengeti Plains. Here we show that modelled populations of Thomson's gazelles can only persist under Serengeti conditions in the face of observed levels of rainfall stochasticity by making adaptive movements to take advantage of ephemeral spatial distributions of food resources. More importantly, our models suggest that Thomson's gazelles in Serengeti require unrestricted access to relatively large areas of grassland (> 1600 km 2 ) to guarantee long-term persistence, particularly when there is positive spatial autocorrelation in resource abundance, as is the case in Serengeti. If this proves to be true for other species and/or other systems, then understanding of complex behavioural responses to spatially and temporally heterogeneous food supplies may be essential to successful conservation of grazing herbivores.